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                  <text>Colorado Division of Parks and Wildlife
September 2013-September 2014
WILDLIFE RESEARCH REPORT
State of:
Cost Center:
Work Package:
Task No.:

Colorado
3420
0660
N/A

Federal Aid
Project No.

N/A

:
:
:
:

Division of Parks and Wildlife
Avian Research
Greater Sage-grouse Conservation
Assessment of Greater Sage-grouse Response to
Pinyon-Juniper Removal in the ParachutePiceance-Roan Population of Northwestern
Colorado

Period Covered: September 1, 2013 – August 31, 2014
Author: B. L. Walker
Personnel: B. Holmes, B. Petch, T. Knowles, B. deVergie
All information in this report is preliminary and subject to further evaluation. Information MAY
NOT BE PUBLISHED OR QUOTED without permission of the author. Manipulation of these data
beyond that contained in this report is discouraged.
ABSTRACT
Greater sage-grouse (Centrocercus urophasianus) in the Parachute-Piceance-Roan (PPR)
region of western Colorado face at least two major potential stressors: projected habitat loss from
energy development and a long-term decline in habitat suitability associated with pinyon-juniper (PJ)
encroachment. PJ removal may be a useful mitigation tool to offset potential habitat losses associated
with energy development. Although PJ removal is commonly used to improve habitat for greater sagegrouse, no studies to date have quantified the timing or magnitude of how birds respond to treatments.
Since 2008, Colorado Parks and Wildlife (CPW) has cooperated with industry and landowner partners
to investigate the effectiveness of PJ removal for restoring sage-grouse habitat in the PPR. In fall
2008, I established nine “survey” study plots, arranged in three groups of three, with each group
consisting of a sagebrush control plot, an untreated PJ control plot, and a PJ treatment plot. Treatments
were completed on the three treatment plots in 2010 and 2011. Pellet surveys over six summers (20092014) indicated that the mean proportion of sample units containing pellets was consistently highest
on sagebrush control plots (range 0.197-0.449 across years), consistently lowest on plots with
encroaching PJ (range 0.007-0.076), and increased starting 1-2 years after treatment, but response was
variable among treatment plots. Twelve transect plots were established in fall 2010 and two more were
added in summer 2011. All 14 transect plots were surveyed for pellets in summer from 2011-2014.
Transect data indicated low mean pellet densities on the four PJ-Control plots over three years (range
across years = 0.00-0.58 pellet piles/km) and on PJ-Treatment plots in the one year prior to treatment
(mean = 0.03 pellet piles/km). Estimates of mean pellet density were substantially higher on four
Sagebrush-Control transect plots over three years (range across years = 11.10 - 27.14 piles/km) and on
one transect plot 4-6 years after treatment (Lower Barnes; range across years = 2.89 - 25.71 piles/km).
There has been no increase in mean pellet density on four treated transect plots within three years after
PJ removal (range across years = 0.00 - 1.04 pellet piles/km). However, estimates of proportion of
sample units with pellets (from survey plots) and of pellet density (from transect plots) also varied

1

�substantially among Sagebrush-Control plots within years and among years within plots, which
suggests there is substantial baseline variation in pellets present, over and above variation in detection
due to observer ability. We completed double-observer sampling on survey plots in 2013 and 2014 to
estimate sample unit-level detectability, and we completed distance sampling on transect plots to
generate distance-detection curves for transects. We established and conducted pre-treatment surveys
on two additional transect plots in summer 2014 (Lower Galloway and Lower Ryan Gulch) in
anticipation of treatments by WPX this fall.

2

�WILDLIFE RESEARCH REPORT
ASSESSMENT OF GREATER SAGE-GROUSE RESPONSE TO PINYON-JUNIPER REMOVAL
IN THE PARACHUTE-PICEANCE-ROAN POPULATION OF NORTHWESTERN COLORADO
BRETT L. WALKER
PROJECT OBJECTIVES
The objective of this study is to measure short-term (&lt; 5 year) responses of greater sage-grouse to
experimental PJ removal using changes in pellet occupancy in a before-after control-treatment
framework.
SEGMENT OBJECTIVES
Objectives of this study from 1 Sept 2013 through 31 August 2014 were to:
1. Conduct pellet surveys on 9 survey plots.
2. Conduct repeat surveys on 9 survey plots to estimate detectability at the sample unit level.
3. Conduct pellet transects on 14 transect plots.
4. Conduct distance sampling for pellets on a subset 4 transect plots.
5. Summarize preliminary results of pellet surveys and transects from 2009-2014.
INTRODUCTION
Large-scale changes to sagebrush ecosystems and historical population declines (Schroeder et
al. 2004) have raised concern about the status and conservation of greater sage-grouse (Centrocercus
urophasianus) and contributed to the recent listing of the species as warranted but precluded under the
Endangered Species Act (DOI 2010). The Parachute-Piceance-Roan (PPR) region holds one of seven
distinct geographic populations of greater sage-grouse in northwestern Colorado. Greater sage-grouse
in the PPR are of conservation concern due to a long-term reduction in habitat suitability caused by
encroachment of pinyon pine (Pinus monophylla) and juniper (Juniperus scopularum and Juniperus
utahensis) into sagebrush and potential impacts from rapidly increasing energy development.
Removal of pinyon-juniper (PJ) from areas with an existing sagebrush understory may help
restore sage-grouse habitat and offset future potential habitat losses from energy development. Pinyonjuniper encroachment into sagebrush over the last 150 years has been identified as a threat to the species’
habitat in the PPR, in Colorado and range-wide (CGSSC 2008). Encroachment is thought to be caused by
fire suppression, reduced fire frequency due to removal of residual grass via livestock grazing, and a
window of climatic conditions suitable for PJ establishment during the late 1800s and early 1990s (Miller
and Rose 1999). Pinyon-juniper removal has been widely implemented in Colorado and range-wide
(CGSSC 2008). However, sage-grouse responses to PJ removal remain poorly studied (Commons et al.
1999), and the timing and magnitude of greater sage-grouse responses following treatment is unknown.
For this reason, it is difficult to judge whether PJ removal can effectively increase available habitat and
offset impacts from energy development.
Since 2008, the Colorado Division of Wildlife, now Colorado Parks and Wildlife (CPW), and
industry and landowner partners have been cooperating on research to assess the value of removing
encroaching PJ as a mitigation strategy in the PPR. The main objective of this study is to measure shortterm (&lt;5 years) responses of greater sage-grouse to experimental PJ removal using changes in pellet
occupancy in a before-after control-treatment framework. This progress report summarizes preliminary
results from summer pellet surveys for the period June 2009 - August 2014.

3

�STUDY AREA
Study plots are within or immediately adjacent to the current occupied range of greater sage-grouse in
the PPR (Fig. 1). Birds in the PPR population inhabit the tops of ridges and plateaus dominated by
mountain big sagebrush (Artemisia tridentata vaseyana) and a mixture of sagebrush and “mountain
shrubs” (e.g., serviceberry, Amelanchier spp.; Gambel oak, Quercus gambelii, snowberry,
Symphoricarpus sp.; antelope bitterbrush, Purshia tridentata, mountain mahogany, Cercocarpus sp.,
etc.). These areas are typically interspersed with patches of aspen (Populus tremuloides) and Douglas-fir
(Pseudotsuga menziesii). Sagebrush and mixed sagebrush-mountain shrub habitats at higher elevation
give way to PJ woodland on lower-elevation ridges that largely precludes use by sage-grouse. Our study
plots are situated along the ecotone where PJ is encroaching upslope into sagebrush and sagebrushmountain shrub habitat.
METHODS
Plot Selection
I used vegetation, topography, marked bird locations, aerial photography, and on-site visits to
identify nine study plots in 2008, 14 additional study plots in 2010 and 2011, and two additional plots in
2014 (Fig. 1). Study plots were selected based on: (1) density of PJ; (2) shrub composition, density, and
height; (3) topography; (4) proximity to areas of known use by greater sage-grouse; and (5) proximity to,
and likelihood of, energy development within five years. All plots had a sagebrush-dominated shrub
layer, typically intermixed with mountain shrubs, and topography similar to habitats used by sage-grouse
in the PPR from 2006-2010 (Apa 2010, Walker 2010). The southeast portion of the PPR population is
experiencing intensive energy development, but there is currently no development within the study plots
and limited development nearby. Two of the plots added in 2011 (Upper and Lower Bar D) were treated
by the Bureau of Land Management in January 2011 and were added opportunistically in our study. An
additional plot (Lower Barnes) added in 2011 was treated by BLM in 2007-2008 and is being surveyed as
a longer-term post-treatment plot for comparison.
Assessing Response to Pinyon-Juniper Removal
I am using a before-after, control-treatment design to compare changes in sage-grouse pellet
occupancy among control and removal plots before and after encroaching PJ is removed. Caution must be
exercised in interpreting results because estimates of pellet occupancy only give an index of frequency of
use during a defined survey period, rather than a measure of abundance, density, habitat quality, or habitat
selection (contra Dahlgren et al. 2006).
I have three levels of treatment: 1) “PJ-Treatment” plots where encroaching PJ is removed, 2)
“PJ-Control” plots where encroaching PJ is not removed, and 3) “Sagebrush-Control” plots with suitable
sagebrush habitat and no PJ. Data from PJ-Treatment plots are used to measure changes in pellet
occupancy before and after PJ removal. PJ-Control plots allow us to measure background changes and
variation in pellet occupancy in areas with encroaching PJ in the absence of treatment. Sagebrush-Control
plots allow us to estimate background changes and variation in pellet occupancy in habitats already
suitable for sage-grouse. Most plots were surveyed for one to three years prior to PJ removal and will be
surveyed for two to five years following removal. I established three study plots per treatment in fall 2008
for a total of nine original “survey” plots. I established 12 additional transect-based study plots in fall
2010 and two more transect-based plots in summer 2011 (Figs. 2-4), for a total of 23 study plots (nine
survey, 14 transect).
Summer Pellet Surveys and Transects

4

�I obtained an index of frequency of sage-grouse use of plots by surveying for pellets during the
summer and estimating pellet occupancy. I used two different sampling techniques for pellet surveys. The
first technique involves surveying a systematic random sample of 30 x 30 m sample units on each
“survey” plot. The second technique involves surveying for pellets within 1.5 m on either side of linear
transects spaced 50 m apart that ran the length of each “transect” study plot.
With both sampling methods, field crews searched for, counted, and removed pellets from each
sample unit (or transect line) within the plot once per year in July-August. For all pellets, field crews
recorded their condition and appearance (crumbly-bleached vs. hard-dry, vs. fresh-wet) to estimate age
and used composition to estimate when during the year they were deposited. Field crews identified those
containing intact insect parts and flower heads as “summer” pellets (April-October) and those containing
only digested sagebrush leaves as “winter” pellets (November-March) (Wallestad et al. 1975). Field
crews recorded single pellets, pellet piles (i.e., day or night roost piles), and cecal droppings separately.
Observers recorded pellets or groups of pellets within 10 cm of each other as one pile, with the constraint
that pellets or groups of pellets &gt; 10 cm apart could not be counted as the same pile.
I opted to sample each plot once per year in July-August such that the “survey period” (i.e., the
period during which birds can deposit pellets) was 12 months. Field crews marked all sample unit centers
and transect lines with aluminum tags, high-visibility stakes and high-visibility flagging to ensure
consistency in sampling locations across years. On survey plots, each observer carefully and thoroughly
surveyed sample units by slowly walking 10 parallel 30-m lines spaced 3 m apart. On transect plots,
observers survey for pellets by walking flagged transect lines and searching within 1.5 m on either side of
the line. Observers recorded anecdotal evidence of occupancy while surveying (e.g., clockers, nests and
eggs, feathers, birds, etc.).
These two survey methods for pellets have the following assumptions: 1) all pellets can be
correctly identified to species; 2) all pellets can be correctly distinguished as either a chick or adult pellet
by size; 3) all pellets deposited during the survey period (during the previous year) can be correctly
distinguished from pellets deposited prior to the survey period by condition and appearance; 4) all pellets
can be correctly distinguished by season (“winter” vs. “summer”) by pellet composition; 5) surveying
does not influence whether or not pellets are present in sampled units (or along transects). To address
assumption 1, I trained observers to distinguish dusky grouse from sage-grouse pellets prior to surveys.
Adult pellets of the two species can be distinguished by composition and smell in any season. Adult-sized
sage-grouse typically consume 13-39% sagebrush throughout the spring and summer and &gt;99%
sagebrush in winter (Wallestad et al. 1975, Schroeder et al. 1999). For this reason, pellets of adult sagegrouse contain sagebrush year-round, and unlike dusky grouse, consistently smell like sage, even after a
year in the field. To address assumption 2, I trained crews to distinguish adult from chick pellets by
length and diameter Quantitative analyses will only include data on adult-size pellets because it may not
be possible to distinguish pellets of dusky grouse vs. sage-grouse chicks due to overlap in chick diets
(both species consume primarily insects and forbs as chicks; Zwickel 1992, Drut et al. 1994). To address
assumption 3, crews differentiated pellets in the field based on condition (bleached and crumbly vs. hard
and dry vs. fresh and green/moist). I am also testing how pellet condition changes with age by placing
piles of fresh test pellets within representative sagebrush habitat in the field and photographing changes in
condition over time. I will test assumption 4 by testing how often observers correctly assess composition
and season for pellets collected from marked birds in different seasons. Assumption 5 may be violated if
surveyors flush birds that then land within another sample unit or along a transect line later surveyed.
However, violation of this assumption is unlikely to meaningfully influence analyses because the number
of pellets birds could deposit before the unit or transect gets surveyed (on the order of minutes or hours) is
miniscule compared to the 12-month survey period.
Pellet Detectability

5

�Detectability of sage-grouse pellets is typically low and may vary among observers, with pellet
condition or appearance, and with distance from observer (Dahlgren et al. 2006). In 2009-2012, I
estimated variation in detectability of individual pellets/pellet piles among observers by having each
observer survey eight test sample units in which I placed piles of fresh pellets of various sizes at random
directions and distances within the sample unit from the sample unit center. Pellets were also placed at
random distances within 1.5 m along two 400-m long test transects in 2011 and 2012.
Field crews also conducted repeat surveys of all survey plots on the same dates in 2013 and 2014
to allow estimation of detectability at the sample-unit level using unreconciled double-observer methods
(Riddle et al. 2010), and conducted distance sampling on all Sagebrush-Control transect plots in 2014 to
allow estimation of pellet distance-detection curves for transect data.
Pinyon-Juniper Removal
All treatments were done by contractors using either a Bobcat with a Fecon head or a Hydro-axe.
Contractors were instructed to remove only pinyon-juniper and to avoid removing sagebrush or mountain
shrubs. A partial treatment was done on the Black Sulphur plot in August 2009 and completed in July
2010, the Upper Galloway plot was treated in November 2010, and the Ryan Gulch plot was treated in
August 2011. Among transect plots, Upper and Lower Bar D plots were treated by BLM in January 2011,
and three plots (Cottonwood, Magnolia South, and Lower Wagonroad) were treated in August-November
2011. Two additional plots added in summer 2014 (Lower Ryan Gulch and Lower Galloway) are adjacent
to two existing study plots (Upper Galloway and Upper Ryan Gulch) and scheduled for treatment by
WPX Energy (in conjunction with CPW and BLM) in fall 2014.
RESULTS AND DISCUSSION
Pellet Surveys and Transects
Pellet survey data from 2009-2014 indicated substantially higher mean seasonal and year-round
pellet detections on Sagebrush-Control survey plots than on PJ-Control plots or on PJ-Treatment plots
prior to treatment (Table 1). Year-round pellet detections increased on the Upper Galloway and Ryan
Gulch plots increased and have remained slightly higher following treatment on both plots since treatment
(Fig. 2, Tables 1). However, there has been no increase in year-round pellets detected on the Black
Sulphur plot within four years following treatment.
Observers conducted repeat surveys of all 9 survey plots on the same dates in 2013 and 2014 to
allow us to estimate detectability of pellets at the sample unit level. Preliminary comparison of resulting
estimates suggests there is substantial variation in observer ability to detect pellets within 30 x 30 m
sample units in both years (Table 2).
Field crews conducted one round of pellet surveys on each of 14 transect plots in July-August in
each year from 2011-2014 (Table 3, Fig. 4). No transect plots showed an increase in the no. pellets
counted per km of transect within two years following treatment, but the mean increased within 3 years
(Fig. 5). However, this increase was driven entirely by an increase on a single plot (Upper Bar D; Fig. 4).
The no. of pellets counted per km of transect was highly variable on the Lower Barnes plot (treated by
BLM in 2007-2008), but generally comparable to Sagebrush-Control plots. No pellets were counted on
pre-treatment transects on the two new plots (Lower Galloway and Lower Ryan Gulch) established in
summer 2014.
Field crews collected distance sampling data on all Sagebrush-Control transect plots in 2014 to
generate pellet distance-detection curves for transect plots. Crews recorded 403 detections, and all pellets
detected were within 3.0 m. These data will be analyzed following additional data collection in 2015.

6

�Taken together, summer pellet data from the pre-treatment phase of the project indicated
substantially greater pellet detections on Sagebrush-Control plots than on either PJ-Control plots or PJTreatment plots prior to treatment.
There has not been a consistent, immediate, or dramatic increase in sage-grouse pellet detections
in treated areas, suggesting that if there is a response by sage-grouse to such habitat treatments, it may
take several years for individuals to colonize and begin using treated areas. Alternatively, sites selected
for removals may still represent marginal habitat, and pellet detections may never increase to levels
comparable to Sagebrush-Control plots. The number of pellets found on Sagebrush-Control plots has also
been highly variable across years, suggesting there substantial natural variation in the number of pellets
deposited (over and above that variation introduced by observer bias). Even so, increases in the number of
pellet detections on 3 of 8 treated plots within 4 years of treatment, combined with consistently low
detections on PJ-Control plots during that same time, suggests that greater sage-grouse do show a positive
response to PJ removal in some areas. Current post-treatment data are insufficient to draw conclusions
about long-term responses of greater sage-grouse to PJ removal.
We anticipate conducting one additional year of pellet surveys (through summer 2015) to ensure
we document grouse response for at least 4-5 years post-treatment. We may continue pellet surveys
beyond 2015 if interest and funding are available, particularly if there is interest in treating PJ-Control
study plots.
LITERATURE CITED
Apa, A. D. 2010. Seasonal habitat use, movements, genetics, and vital rates in the Parachute-PiceanceRoan Population of greater sage-grouse. Unpublished progress report. Colorado Division of
Wildlife. Fort Collins, USA.
Becker, E. F. 1991. A terrestrial furbearer estimator based on probability sampling. Journal of Wildlife
Management 55: 730-737.
Becker, E. F., Golden, H. N., and Gardner, C. L. 2004. Using probability sampling of animal tracks in
snow to estimate population size. In Thompson, W. L., ed., Sampling rare or elusive species:
concepts and techniques for estimating population parameters. Island Press, Washington, D. C.,
USA, pp. 248-270.
Becker, E. F., M. A. Spindler, and T. O. Osborne. 1998. A population estimator based on network
sampling of tracks in the snow. Journal of Wildlife Management 62:968-977.
Canfield, R. H. 1941. Application of the line interception method in sampling range vegetation. Journal of
Forestry 39:388–394.
Colorado Greater Sage-Grouse Steering Committee (CGSSC). 2008. Colorado greater sage-grouse
conservation plan. Colorado Division of Wildlife, Denver, USA.
Commons, M. L., R. K. Baydack, and C. E. Braun. 1999. Sage grouse response to pinyon-juniper
management. Pages 238-239 in S. B. Monsen and R. Stevens, compilers. Proceedings: ecology
and management of pinyon-juniper communities. RMRS-P-9. United States Department of
Agriculture, Forest Service, Fort Collins, Colo. USA.
Dahlgren, D. K., R. Chi, and T. A. Messmer. 2006. Greater sage-grouse response to sagebrush
management in Utah. Wildlife Society Bulletin 34:975-985.
Department of the Interior (DOI). 2010. 12-month finding for petitions to list the greater sage-grouse as
threatened or endangered. Federal Register 75(55): 13910-14014.
Drut, M. S., W. H. Pyle, and J. A. Crawford. 1994. Technical note: Diets and food selection of Sage
Grouse chicks in Oregon. Journal of Range Management 47:90-93.
Miller, R. F. and J. A. Rose.1999. Fire history and western juniper encroachment in sagebrush steppe.
Journal of Range Management 52:550-559.

7

�Schroeder, M. A., C. L. Aldridge, A. D. Apa, J. R. Bohne, C. E. Braun, S. D. Bunnell, J. W. Connelly, P.
A. Deibert, S. C. Gardner, M. A. Hilliard, G. D. Kobriger, C. W. Mccarthy. 2004. Distribution of
Sage-grouse in North America. Condor 106:363-376.
Schroeder, M. A., J. R. Young, and C. E. Braun. 1999. Sage-grouse (Centrocercus urophasianus).
Account 425 in A. Poole and F. Gill, editors. The birds of North America. The Academy of
Natural Sciences, Philadelphia, Pa. USA.
Walker, B. L. 2010. Greater sage-grouse research in the Parachute-Piceance-Roan region of western
Colorado: multi-scale habitat selection and seasonal habitat mapping. Unpublished interim
progress report. Colorado Division of Wildlife. Fort Collins, USA.
Wallestad, R., J. G. Peterson, and R. L. Eng. 1975. Foods of adult sage grouse in central Montana. Journal
of Wildlife Management 39:628-630.
Zwickel, F. C. 1992. Blue Grouse (Dendragapus obscurus). Account 15 in A. Poole, P. Stettenheim, and
F. Gill, editors. The birds of North America. The Academy of Natural Sciences, Philadelphia, Pa.
USA.

8

�Table 1. Preliminary estimates of proportion of sample units ± SE containing greater sage-grouse pellets (from any season) on survey plots in the ParachutePiceance-Roan population of western Colorado, USA from 2009-2014. Values presented do not account for sample-unit level variation in pellet detectability.
PJ - Treatment Plotsa

PJ - Control (No Treatment) Plots

Sagebrush - Control Plots

1-Upper 2-Black
3-Ryan
MEAN
1-Dry
3-Stake
MEAN
1-Dry
2-Canyon
3-Black
MEAN
Galloway Sulphur
Gulch
± SE
Ryan
2-Eureka
Springs
± SE
Gulch
Creek
Cabin
± SE
b
b
b
c
Year
n = 49
n = 38
n = 48
N=3
n = 35
n = 50
n = 38
N=3
n = 54
n = 41
n = 42
N=3
d
2009
0.082
0.000
0.146
0.076
0.000
0.040
0.105
0.048
0.444
0.293
0.452
0.397
± 0.039 ± 0.000
± 0.051 ± 0.042
± 0.000
± 0.033
± 0.05
± 0.031
± 0.068
± 0.071
± 0.077 ± 0.052
2010
0.000
0.104
0.052
0.029
0.040
0.026
0.032
0.721
0.293
0.333
0.449
0.026
± 0.000 ± 0.026
± 0.044 ± 0.052
± 0.028
± 0.028
± 0.026
± 0.004
± 0.068
± 0.071
± 0.073 ± 0.136
2011
0.063
0.000
0.020
0.000
0.007
0.349
0.171
0.071
0.197
0.082
0.000
0.041
± 0.035 ± 0.041
± 0.000
± 0.020
± 0.000
± 0.007
± 0.073
± 0.059
± 0.040 ± 0.081
± 0.039 ± 0.000
2012
0.029
0.040
0.000
0.023
0.605
0.341
0.262
0.403
0.245
0.026
0.188
0.153
± 0.028
± 0.028
± 0.000
± 0.012
± 0.075
± 0.074
± 0.068 ± 0.104
± 0.061 ± 0.026
± 0.056 ± 0.065
2013
0.000
0.100
0.000
0.033
0.605
0.220
0.381
0.402
0.082
0.000
0.188
0.090
± 0.000
± 0.042
± 0.000
± 0.033
± 0.075
± 0.065
± 0.075 ± 0.112
± 0.039 ± 0.000
± 0.056 ± 0.054
2014
0.000
0.000
0.026
0.009
0.535
0.146
0.381
0.354
0.082
0.000
0.333
0.138
± 0.000
± 0.000
± 0.026
± 0.009
± 0.076
± 0.055
± 0.075 ± 0.113
± 0.039 ± 0.000
± 0.068 ± 0.100
a
Values in bold represent post-treatment data. Numbers before plot names refer to set (e.g., Set 1 = Upper Galloway, Dry Ryan, Dry Gulch, etc.).
b
n refers to no. of 30 x 30 m units sampled within the study plot.
c
N refers to no. of study plots contributing to the mean. Means in bold are for post-treatment plots, otherwise values represent pre-treatment data.
d
p = proportion of sample units per plots in which greater sage-grouse pellets were detected. Subscripts refer to year.

9

�Table 2. Observer variation in estimates of proportion of sample units with greater sage-grouse pellets (from any season) on survey plots from repeat surveys in the
Parachute-Piceance-Roan population of western Colorado, USA in 2013 and 2014. Values presented do not account for variation in sample-unit level detectability
of pellets. Surveys of each sample unit were conducted by a different observer within 20-30 minutes of each other on the same date.
PJ - Treatment Plots (post-treatment)
PJ - Control (No Treatment) Plots
Sagebrush - Control Plots
1-Upper
Galloway
n = 49a
pBc

3-Ryan
Gulch
n = 48a

MEAN ±
SE
N = 3b

0.000
± 0.000
0.000
± 0.000

0.188
± 0.056
0.167
± 0.054

0.090
± 0.054
0.130
± 0.067

1-Dry
Ryan
n = 35

3-Stake
Springs
n = 38

MEAN ±
SE
N=3

0.000
± 0.000
0.000
± 0.000

0.033
± 0.033
0.013
± 0.013

1-Dry
Gulch
n = 43

2-Canyon
Creek
n = 41

3-Black
Cabin
n = 42

MEAN ±
SE
N=3

0.220
± 0.065
0.195
± 0.062

0.381
± 0.075
0.238
± 0.066

0.402
± 0.112
0.299
± 0.084

0.184
0.000
0.250
0.145
0.029
0.026
0.052
0.488
± 0.055 ± 0.000 ± 0.063
± 0.075
± 0.028
± 0.026
± 0.024
± 0.076
pAc
0.082
0.000
0.333
0.138
0.000
0.026
0.009
0.535
± 0.039 ± 0.000 ± 0.068
± 0.100
± 0.000
± 0.026
± 0.009
± 0.076
a
n refers to the number of sample units per study plot.
b
N refers to the no. of study plots contributing to the mean.
c
p = proportion of sample units in which greater sage-grouse pellets were detected. Subscript refers observer (A or B).

0.098
± 0.046
0.146
± 0.055

0.548
± 0.077
0.381
± 0.075

0.378
± 0.141
0.354
± 0.113

pBc

0.000
± 0.000
0.000
± 0.000

2-Eureka
n = 50
2013
0.100
± 0.042
0.040
± 0.028
2014
0.100
± 0.042
0.000
± 0.000

0.605
± 0.075
0.465
± 0.076

pAc

0.082
± 0.039
0.224
± 0.06

2-Black
Sulphur
n = 38a

10

�Table 3. Preliminary estimates of the density of greater sage-grouse pellets (from any season) encountered on transect plots in the Parachute-Piceance-Roan
population of western Colorado, USA from 2011-2014. Values presented do not account for variation in pellet detectability with distance.
PJ - Treatment Plotsa
PJ - Control (No Treatment) Plots
Sagebrush - Control Plots
Lower
Upper
Bar
Upper WagonMagnolia Cotton- Wagon- Lower Upper
Lower
Magnolia
WagonD
Magnolia Barnes
road
Bar D
South
wood
road
Bar D Bar D
Barnes
North
Sprague
road
Split
Control
Control Control Control
x2011b
0.10
0.00
0.00
0.00
0.00
0.00
0.00
18.82
13.51
6.46
5.61
0.00
0.00
13.78
c
X2011
0.03 ± 0.03
0.00 ± 0.00
11.10 ± 3.84
0.00 ± 0.00
x2012
X2012

2.02
0.00
0.31
0.00
46.03
7.94
31.63
22.96
0.00
0.00
0.63
25.51
0.58 ± 0.42
27.14 ± 7.97
0.75 ± 0.54
x2013
0.00
0.00
0.00
0.00
18.53
1.01
11.56
60.03
0.31
0.00
0.00
0.52
0.00
2.89
X2013
0.00 ± 0.00
22.79 ± 12.93
0.17 ± 0.11
x2014
0.00
0.00
0.10
0.00
19.41
4.22
15.31
33.16
2.14
0.00
0.00
0.10
8.96
3.91
X2014
0.00 ± 0.00
18.03 ± 5.98
2.24 ± 1.73
d
km
9.80
7.52
7.64
9.58
6.36
5.88
6.44
12.52
9.80
4.68
6.80
5.92
5.88
5.88
(7.70)
(13.40)
a
Post-treatment data are in bold. Treatments on Lower Bar D and Upper Bar D occurred in Jan 2011. Data for Lower Bar D and Upper Bar D from 2011-2014
represent ~6, 18, 30, and 42 mo post-treatment, respectively. Treatments on Lower Barnes were started by the Bureau of Land Management in summer 2007
and completed in summer 2008. Post-treatment data on Lower Barnes were collected opportunistically and are presented for comparison only.
b
x = no. pellet piles detected per km of transect.
c
X = mean no. pellet piles detected per km of transect across study plots within each treatment. Values for Lower Barnes are presented separately. The 2011
mean for Lower Bar D and Upper Bar D in 2011 represents post-treatment data and is presented separately from the 2011 mean for other PJ – Treatment plots
that had not yet been treated (Magnolia South, Cottonwood, Lower Wagonroad).
d
Total km of transect line surveyed per plot. Portions of transects on Cottonwood and Sprague unsuitable as greater sage-grouse habitat were eliminated in
2012, values in parentheses indicate km surveyed in 2011 only.
2.86

0.27

11

�Table 4. Timeline for research on greater sage-grouse response to pinyon-juniper removal in the ParachutePiceance-Roan population, western Colorado, 2008-2015.
Task
Identification of plots for PJ removal
Remove encroaching PJ on survey treatment plots (2009-2011)
Remove encroaching PJ on transect treatment plots (2011)
Remove encroaching PJ on new transect treatment plots (2014)
Pellet surveys (annually)
Prepare cumulative report (annually)
Prepare cumulative final report

12

Initiation
Summer 2008
Fall 2009
Fall 2011
Fall 2014
25 June
1 Sep
1 Aug 2015

Completion
COMPLETED
COMPLETED
COMPLETED
Fall 2014
31 Aug
30 Sep
30 Sep 2015

�Fig. 1. Map of study plot locations from 2009-2014 for the pinyon-juniper removal experiment and greater sage-grouse occupied
range boundary (as of 2012) in the northern portion of the Parachute-Piceance-Roan population of western Colorado, USA.

13

�Fig. 2. Trends in the proportion of sample units detected with greater sage-grouse pellets for individual survey plots (color-coded by
treatment) in the Parachute-Piceance-Roan population of western Colorado, USA, 2009-2014. Values presented do not account for
variation in sample-unit level detectability of pellets.

14

�Fig. 3. Trends in the mean proportion of sample units detected with greater sage-grouse pellets across survey plots (color-coded by
treatment) in the Parachute-Piceance-Roan population of western Colorado, USA, 2009-2014. Values presented do not account for
variation in sample-unit level detectability of pellets.

15

�Fig. 4. Trends in the no. of greater sage-grouse pellets counted per km of transect for individual transect plots (color-coded by treatment)
in the Parachute-Piceance-Roan population of western Colorado, USA, 2011-2014. Values presented do not account for variation in
detectability of pellets with distance from transect.

16

�Fig. 5. Trends in the mean no. of greater sage-grouse pellets counted per km of transect across transect plots within treatments (colorcoded by treatment) in the Parachute-Piceance-Roan population of western Colorado, USA, 2011-2014. Values presented do not account
for variation in detectability of pellets with distance from transect.

17

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                  <text>Colorado Division of Parks and Wildlife
September 2014-September 2015
WILDLIFE RESEARCH REPORT
State of:
Cost Center:
Work Package:
Task No.:

Colorado
3420
0660
N/A

Federal Aid
Project No.

N/A

:
:
:
:

Division of Parks and Wildlife
Avian Research
Greater Sage-grouse Conservation
Assessment of Greater Sage-grouse Response to
Pinyon-Juniper Removal in the ParachutePiceance-Roan Population of Northwestern
Colorado

Period Covered: September 1, 2014 – August 31, 2015
Author: B. L. Walker
Personnel: B. Holmes, B. Petch, T. Knowles, B. deVergie
All information in this report is preliminary and subject to further evaluation. Information MAY
NOT BE PUBLISHED OR QUOTED without permission of the author. Manipulation of these data
beyond that contained in this report is discouraged.
EXTENDED ABSTRACT
Greater sage-grouse (Centrocercus urophasianus) in the Parachute-Piceance-Roan (PPR) region of
western Colorado face at least two major potential stressors: projected habitat loss from energy
development and a long-term decline in habitat suitability associated with pinyon-juniper (PJ)
encroachment. Pinyon-juniper removal may be a useful mitigation tool to offset potential habitat
losses associated with energy development. Although pinyon-juniper removal is commonly used to
improve habitat for greater sage-grouse, no studies to date have quantified the timing or magnitude of
how birds respond to treatments. Since 2008, Colorado Parks and Wildlife (CPW) has cooperated with
industry and landowner partners to investigate the effectiveness of pinyon-juniper removal for
restoring sage-grouse habitat in the PPR. In fall 2008, I established nine “survey” study plots, arranged
in three groups of three, with each group consisting of a Sagebrush-Control plot, an untreated PJControl plot, and a PJ-treatment plot. Treatments were completed on three survey plots in 2010 and
2011. Pellet surveys in summer from 2009-2015 indicated that the mean proportion of sample units
containing pellets was consistently highest on sagebrush control plots (range 0.197-0.449 across
years), consistently lowest on plots with encroaching pinyon-juniper (range 0.007-0.048), and
increased on 2 of 3 treated survey plots (Ryan Gulch and Upper Galloway) within 1-2 years after
treatment. Fourteen transect plots were established in fall 2010 and summer 2011, and two were
established in summer 2014. Transect plots were surveyed for pellets in summer though 2015. As
expected, estimated mean pellet piles/km were low on the four PJ-Control plots for the duration of the
study (range across years = 0.00-0.58 pellets/km) and on PJ-Treatment plots prior to treatment (mean
= 0.03 pellet piles/km). Mean pellet piles/km were consistently higher on all four Sagebrush-Control
transect plots through 2014 (range across years = 11.10 - 27.14 pellet piles/km), but declined
precipitously on 3 of 4 Sagebrush-Control plots in 2015 (Magnolia Control, Upper Barnes Control,
and Wagonroad Control). Pellet piles/km were also high on the Lower Barnes transect plot 4-5 years
post-treatment (13.78-25.71 piles/km), but declined 6-8 years post-treatment (0.00-3.91 pellet

1

�piles/km). Mean pellet piles/km has generally remained low on treated transect plots for four years
after pinyon-juniper removal (range across years = 0.00 - 2.86 pellet piles/km) with the exception of
Upper Bar D in 2014 (8.96 pellet piles/km). Estimates of proportion of sample units with pellets (from
survey plots) and of pellet piles/km (from transect plots) also varied substantially among SagebrushControl plots within years and among years within plots, which suggests there is substantial variation
in pellets deposited within suitable habitat (i.e., over and above variation in counts due to observer
bias). We completed double-observer sampling on survey plots in 2013, 2014, and 2015 to estimate
sample unit-level detectability, and we completed distance sampling on transect plots in 2014 and
2015 to generate distance-detection curves, and those analyses are in progress. We established and
conducted pre- and post-treatment surveys on two additional transect plots (Lower Galloway and
Lower Ryan Gulch) in summer 2014 and 2015. To date, response to pinyon-juniper removal has been
inconsistent, with pellet counts increasing on only two of eight plots within 4-5 years post-treatment.
However, a recent experimental treatment suggests that sage-grouse may respond more strongly to
combined pinyon-juniper and serviceberry removal. We are developing a proposal to expand the
current pinyon-juniper removal project to include serviceberry treatments on existing study plots
starting in fall 2017.

2

�COLORADO PARKS AND WILDLIFE RESEARCH REPORT
ASSESSMENT OF GREATER SAGE-GROUSE RESPONSE TO PINYON-JUNIPER REMOVAL
IN THE PARACHUTE-PICEANCE-ROAN POPULATION OF NORTHWESTERN COLORADO
BRETT L. WALKER

INTRODUCTION
Large-scale changes to sagebrush ecosystems and historical population declines (Schroeder et
al. 2004) have raised concern about the status and conservation of greater sage-grouse (Centrocercus
urophasianus) and contributed to the recent listing of the species as warranted but precluded under the
Endangered Species Act (DOI 2010). The Parachute-Piceance-Roan (PPR) region holds one of seven
distinct geographic populations of greater sage-grouse in northwestern Colorado. Greater sage-grouse
in the PPR are of conservation concern due to a long-term reduction in habitat suitability caused by
encroachment of pinyon pine (Pinus monophylla) and juniper (Juniperus scopularum and Juniperus
utahensis) into sagebrush and potential impacts from rapidly increasing energy development.
Removal of pinyon-juniper from areas with an existing sagebrush understory may help restore
sage-grouse habitat and offset future potential habitat losses from energy development. Pinyon-juniper
encroachment into sagebrush over the last 150 years has been identified as a threat to the species’ habitat
in the PPR, in Colorado and range-wide (CGSSC 2008). Encroachment is thought to be caused by fire
suppression, reduced fire frequency due to removal of residual grass via livestock grazing, and a window
of climatic conditions suitable for pinyon-juniper establishment during the late 1800s and early 1990s
(Miller and Rose 1999). Pinyon-juniper removal has been widely implemented in Colorado and rangewide (CGSSC 2008). However, sage-grouse responses to pinyon-juniper removal remain poorly studied
(Commons et al. 1999), and the timing and magnitude of greater sage-grouse responses following
treatment is unknown. For this reason, it is difficult to judge whether pinyon-juniper removal can
effectively increase available habitat and offset impacts from energy development.
Since 2008, the Colorado Division of Wildlife, now Colorado Parks and Wildlife (CPW), and
industry and landowner partners have been cooperating on research to assess the value of removing
encroaching pinyon-juniper as a mitigation strategy in the PPR. The main objective of this study is to
measure short-term (&lt;5 years) responses of greater sage-grouse to experimental pinyon-juniper removal
using changes in pellet occupancy in a before-after control-treatment framework. This progress report
summarizes preliminary results from summer pellet surveys for the period June 2009 - August 2015.
STUDY AREA
Study plots are within or immediately adjacent to the current occupied range of greater sage-grouse in
the PPR (Fig. 1). Birds in the PPR population inhabit the tops of ridges and plateaus dominated by
mountain big sagebrush (Artemisia tridentata vaseyana) and a mixture of sagebrush and “mountain
shrubs” (e.g., serviceberry, Amelanchier spp.; Gambel oak, Quercus gambelii, snowberry,
Symphoricarpus sp.; antelope bitterbrush, Purshia tridentata, mountain mahogany, Cercocarpus sp.,
etc.). These areas are typically interspersed with patches of aspen (Populus tremuloides) or conifers.
Sagebrush and mixed sagebrush-mountain shrub habitats at higher elevation give way to pinyon-juniper
woodland on lower-elevation ridges that largely precludes use by sage-grouse. Our study plots are
situated along the ecotone where pinyon-juniper is encroaching upslope into sagebrush and mixed
sagebrush-mountain shrub habitat.

3

�METHODS
Plot Selection
I used vegetation, topography, marked bird locations, aerial photography, and on-site visits to
identify nine study plots in 2008, 14 additional study plots in 2010 and 2011, and two additional plots in
2014 (Fig. 1). Study plots were selected based on: (1) density of pinyon-juniper; (2) shrub composition,
density, and height; (3) topography; (4) proximity to areas of known use by greater sage-grouse; and (5)
proximity to, and likelihood of, energy development within five years. All plots had a sagebrushdominated shrub layer, typically intermixed with mountain shrubs, and topography similar to habitats
used by sage-grouse in the PPR from 2006-2010 (Apa 2010, Walker et al. 2015). The southeast portion of
the PPR population is experiencing intensive energy development, but there is currently no development
within the study plots and limited development nearby. Two of the plots added in 2011 (Upper and Lower
Bar D) were treated by the Bureau of Land Management in January 2011 and were added
opportunistically to our study. An additional plot (Lower Barnes) added in 2011 was treated by BLM in
2007-2008 and has been included in our study as a longer-term post-treatment plot for comparison.
Assessing Response to Pinyon-Juniper Removal
I used a before-after, control-treatment design to compare changes in sage-grouse pellet
occupancy among control and removal plots before and after encroaching pinyon-juniper is removed.
Caution must be exercised in interpreting results because estimates of pellet occupancy only give an index
of frequency of use during a defined survey period, rather than a measure of abundance, density, habitat
quality, or habitat selection (contra Dahlgren et al. 2006). There are three levels of treatment: 1) “PJTreatment” plots where encroaching pinyon-juniper is removed, 2) “PJ-Control” plots where encroaching
pinyon-juniper is not removed, and 3) “Sagebrush-Control” plots with suitable sagebrush habitat and no
pinyon-juniper encroachment. Data from PJ-Treatment plots are used to measure changes in pellet
occupancy before and after removal. PJ-Control plots allow us to measure background changes and
variation in pellet occupancy in areas with encroaching pinyon-juniper in the absence of treatment.
Sagebrush-Control plots allow us to estimate background changes and variation in pellet occupancy in
habitats already suitable for sage-grouse. Most plots were surveyed for one to three years prior to pinyonjuniper removal and will be surveyed for two to five years following removal. I established three study
plots per treatment in fall 2008 for a total of nine original “survey” plots. I established 12 additional
transect-based study plots in fall 2010 and two more transect-based plots in summer 2011 (Figs. 2-4), for
a total of 23 study plots (nine survey, 14 transect).
Summer Pellet Surveys and Transects
I obtained an index of frequency of sage-grouse use of plots by surveying for pellets during the
summer and estimating pellet occupancy. I used two different sampling techniques for pellet surveys. The
first technique involves surveying a systematic random sample of 30 x 30 m sample units on each
“survey” plot. The second technique involves surveying for pellets within 1.5 m on either side of linear
transects spaced 50 m apart that ran the length of each “transect” study plot.
With both sampling methods, field crews searched for, counted, and removed pellets from each
sample unit (or transect line) within the plot once per year in July-August. For all pellets, field crews
recorded their condition and appearance (crumbly-bleached vs. hard-dry, vs. fresh-wet) to estimate age
and used composition to estimate when during the year they were deposited. Field crews identified those
containing intact insect parts and flower heads as “summer” pellets (April-October) and those containing
only digested sagebrush leaves as “winter” pellets (November-March) (Wallestad et al. 1975). Field
crews recorded single pellets, pellet piles (i.e., day or night roost piles), and cecal droppings separately.
Observers recorded pellets or groups of pellets within 10 cm of each other as one pile, with the constraint
that pellets or groups of pellets &gt; 10 cm apart could not be counted as the same pile.

4

�I opted to sample each plot once per year in July-August such that the “survey period” (i.e., the
period during which birds can deposit pellets) was 12 months. Field crews marked all sample unit centers
and transect lines with aluminum tags, high-visibility stakes and high-visibility flagging to ensure
consistency in sampling locations across years. On survey plots, each observer carefully and thoroughly
surveyed sample units by slowly walking 10 parallel 30-m lines spaced 3 m apart. On transect plots,
observers survey for pellets by walking flagged transect lines and searching within 1.5 m on either side of
the line. Observers recorded anecdotal evidence of occupancy while surveying (e.g., clockers, nests and
eggs, feathers, birds, etc.).
These methods have the following assumptions: 1) all pellets are correctly identified to species; 2)
all pellets are correctly distinguished as either a chick or adult by size; 3) all pellets deposited during the
survey period (during the previous year) are correctly distinguished from pellets deposited prior to the
survey period by condition and appearance; 4) all pellets are correctly distinguished by season (“winter”
vs. “summer”) by pellet composition; 5) surveying does not influence whether or not pellets are present in
sampled units (or along transects). To address assumption 1, I trained observers to distinguish dusky
grouse from sage-grouse pellets prior to surveys. Adult pellets of the two species can be distinguished by
composition and smell in any season. Adult-sized sage-grouse typically consume 13-39% sagebrush
throughout the spring and summer and &gt;99% sagebrush in winter (Wallestad et al. 1975, Schroeder et al.
1999). For this reason, pellets of adult sage-grouse contain sagebrush year-round, and unlike dusky
grouse, consistently smell like sage, even after a year in the field. To address assumption 2, I trained
crews to distinguish adult from chick pellets by length and diameter. Analyses will only include data on
adult-size pellets because it may not be possible to distinguish pellets of dusky grouse vs. sage-grouse
chicks due to overlap in chick diets (both species consume primarily insects and forbs as chicks; Zwickel
1992, Drut et al. 1994). To address assumption 3, crews differentiated pellets in the field based on
condition (bleached and crumbly vs. hard and dry vs. fresh and green/moist). I am also testing how pellet
condition changes with age by placing piles of fresh test pellets within representative sagebrush habitat in
the field and photographing changes in condition over time. I will test assumption 4 by testing how often
observers correctly assess composition and season for pellets collected from marked birds in different
seasons. Assumption 5 may be violated if surveyors flush birds that then land within another sample unit
or along a transect line later surveyed. However, violation of this assumption is unlikely to meaningfully
influence analyses because the number of pellets birds could deposit before the unit or transect gets
surveyed (on the order of minutes or hours) is insignificant compared to the 12-month survey period.
Pellet Detectability
Detectability of sage-grouse pellets is typically low and may vary among observers, with pellet
condition or appearance, and with distance from observer (Dahlgren et al. 2006). In 2009-2012, I
estimated variation in detectability of individual pellets/pellet piles among observers by having each
observer survey eight test sample units in which I placed piles of fresh pellets of various sizes at random
directions and distances within the sample unit from the sample unit center. Pellets were also placed at
random distances within 1.5 m along two 400-m long test transects in 2011 and 2012.
Field crews also conducted repeat surveys of all survey plots on the same dates in 2013, 2014,
and 2015 to allow estimation of detectability at the sample-unit level using unreconciled double-observer
methods (Riddle et al. 2010), and conducted distance sampling on all Sagebrush-Control transect plots in
2014 and 2015 to allow estimation of pellet distance-detection curves for transect data.
Pinyon-Juniper Removal
All treatments were done by contractors using either a Bobcat with a Fecon head or a Hydro-axe.
Contractors were instructed to remove only pinyon-juniper and to avoid removing sagebrush or mountain
shrubs. Treatment started on the Black Sulphur plot in August 2009 and was completed in July 2010, the
Upper Galloway plot was treated in November 2010, and the Ryan Gulch plot was treated in August

5

�2011. Among transect plots, Upper and Lower Bar D plots were treated by BLM in January 2011, and
three plots (Cottonwood, Magnolia South, and Lower Wagonroad) were treated in August-November
2011. Two additional plots added in summer 2014 (Lower Ryan Gulch and Lower Galloway) are adjacent
to two existing study plots (Upper Galloway and Upper Ryan Gulch) and were treated by WPX Energy
(in conjunction with CPW and BLM) in fall 2014.
RESULTS AND DISCUSSION
Pellet Surveys and Transects
Pellet survey data from 2009-2015 indicated substantially higher mean seasonal and year-round
pellet detections on Sagebrush-Control survey plots than on PJ-Control plots or on PJ-Treatment plots
prior to treatment (Table 1). Year-round pellet detections increased on the Upper Galloway and Ryan
Gulch plots and have remained higher for 5 years following treatment (Fig. 2, Tables 1). However, no
increase in pellets was detected on the Black Sulphur plot within 5 years following treatment. Preliminary
comparison of repeated-count data suggests substantial variation in observer ability to detect pellets
within 30 x 30 m sample units in all three years (Table 2). Analysis of repeated-count data are in progress.
Field crews conducted one round of pellet surveys on each of 14 transect plots in July-August in
each year from 2011-2015 (Table 3, Fig. 4). No transect plots showed any substantive increase in the no.
pellet piles/km within four years following treatment with the exception of a temporary increase on a
single plot in a single year (Upper Bar D in 2014). The no. of pellets counted per km of transect on the
Lower Barnes plot (treated by BLM in 2007-2008) was comparable to Sagebrush-Control plots in 20112012, but decreased to levels comparable to those of PJ-Treatment plots from 2013-2015 (Table 3, Fig.
4). No pellets were counted on pre-treatment transects on the two new plots (Lower Galloway and Lower
Ryan Gulch) established in 2014, but a handful pellets were detected post-treatment in 2015. Field crews
collected distance sampling data on all Sagebrush-Control transect plots in 2014 and 2015 to generate
pellet distance-detection curves for transect plots. Crews recorded 403 detections in 2014 and 273
detections in 2015. All pellets detected were within 2.85 m of the transect line. These data are currently
being analyzed using program DISTANCE to generate distance-detection curves.
Overall, summer pellet data indicated substantially greater pellet detections on Sagebrush-Control
plots than on either PJ-Treatment plots prior to treatment or on PJ-Control plots as expected, but they also
demonstrate substantial variation in pellet counts within and among Sagebrush-Control plots over time.
An increase in sage-grouse pellet detections has only been observed on 2 of 8 treated plots within
4-5 years post-treatment. It is possible that populations take longer to respond to such treatments if few
yearlings are available to colonize new habitat. However, our study was long enough to capture several
years of high abundance. Alternatively, there may be unmeasured factors that hinder the response on
some sites (e.g., too much surrounding forest). Also, sites selected for removals were generally on the
edge of occupied range and may simply represent marginal habitat that gets reduced or occasional use. In
that case, pellet detections may never increase to levels comparable to Sagebrush-Control plots. The
number of pellets found on Sagebrush-Control plots has also been highly variable across years (this was
especially evident in 2015) which suggests that there is substantial variation in the number of pellets
deposited within any given area of suitable habitat as birds move around from year to year (over and
above variation in counts due to observer bias). Such variation may be because our study sites are within
a dynamic landscape and changes in the suitability of areas surrounding our study plots (e.g., due to
energy development or other habitat treatments) may cause birds to shift which habitats they use from
year to year. Such variation could also be attributable to birds moving around within our study plots
enough that current sampling intensity is insufficient to consistently detect within-plot use.
Although our data suggest a positive response of greater sage-grouse on at least two study plots,
pinyon-juniper removal does not appear to be a consistently reliable mitigation option in the PPR. For that

6

�reason, we need to consider other habitat treatment options. An experimental project by Conoco-Phillips
from 2013-2015 looked at changes in sage-grouse pellet detections and micro-site vegetation in response
to combined pinyon-juniper and serviceberry treatments. Based on the apparent initial success of that
project, we are now considering incorporating serviceberry treatments into our existing project. We would
either conduct serviceberry treatments on our existing PJ-Treatment plots or conduct combined pinyonjuniper and serviceberry treatments on existing PJ-Control plots and continue pellet surveys to document
initial response to such treatments.
LITERATURE CITED
Apa, A. D. 2010. Seasonal habitat use, movements, genetics, and vital rates in the Parachute-PiceanceRoan Population of greater sage-grouse. Unpublished progress report. Colorado Division of
Wildlife. Fort Collins, USA.
Becker, E. F. 1991. A terrestrial furbearer estimator based on probability sampling. Journal of Wildlife
Management 55: 730-737.
Becker, E. F., Golden, H. N., and Gardner, C. L. 2004. Using probability sampling of animal tracks in
snow to estimate population size. In Thompson, W. L., ed., Sampling rare or elusive species:
concepts and techniques for estimating population parameters. Island Press, Washington, D. C.,
USA, pp. 248-270.
Becker, E. F., M. A. Spindler, and T. O. Osborne. 1998. A population estimator based on network
sampling of tracks in the snow. Journal of Wildlife Management 62:968-977.
Canfield, R. H. 1941. Application of the line interception method in sampling range vegetation. Journal of
Forestry 39:388–394.
Colorado Greater Sage-Grouse Steering Committee (CGSSC). 2008. Colorado greater sage-grouse
conservation plan. Colorado Division of Wildlife, Denver, USA.
Commons, M. L., R. K. Baydack, and C. E. Braun. 1999. Sage grouse response to pinyon-juniper
management. Pages 238-239 in S. B. Monsen and R. Stevens, compilers. Proceedings: ecology
and management of pinyon-juniper communities. RMRS-P-9. United States Department of
Agriculture, Forest Service, Fort Collins, Colo. USA.
Dahlgren, D. K., R. Chi, and T. A. Messmer. 2006. Greater sage-grouse response to sagebrush
management in Utah. Wildlife Society Bulletin 34:975-985.
Department of the Interior (DOI). 2010. 12-month finding for petitions to list the greater sage-grouse as
threatened or endangered. Federal Register 75(55): 13910-14014.
Drut, M. S., W. H. Pyle, and J. A. Crawford. 1994. Technical note: Diets and food selection of Sage
Grouse chicks in Oregon. Journal of Range Management 47:90-93.
Miller, R. F. and J. A. Rose.1999. Fire history and western juniper encroachment in sagebrush steppe.
Journal of Range Management 52:550-559.
Schroeder, M. A., C. L. Aldridge, A. D. Apa, J. R. Bohne, C. E. Braun, S. D. Bunnell, J. W. Connelly, P.
A. Deibert, S. C. Gardner, M. A. Hilliard, G. D. Kobriger, C. W. Mccarthy. 2004. Distribution of
Sage-grouse in North America. Condor 106:363-376.
Schroeder, M. A., J. R. Young, and C. E. Braun. 1999. Sage-grouse (Centrocercus urophasianus).
Account 425 in A. Poole and F. Gill, editors. The birds of North America. The Academy of
Natural Sciences, Philadelphia, Pa. USA.
Walker, B. L. 2010. Greater sage-grouse research in the Parachute-Piceance-Roan region of western
Colorado: multi-scale habitat selection and seasonal habitat mapping. Unpublished interim
progress report. Colorado Division of Wildlife. Fort Collins, USA.
Wallestad, R., J. G. Peterson, and R. L. Eng. 1975. Foods of adult sage grouse in central Montana. Journal
of Wildlife Management 39:628-630.
Zwickel, F. C. 1992. Blue Grouse (Dendragapus obscurus). Account 15 in A. Poole, P. Stettenheim, and
F. Gill, editors. The birds of North America. The Academy of Natural Sciences, Philadelphia, Pa.
USA.

7

�Table 1. Preliminary estimates of proportion of sample units ± SE containing greater sage-grouse pellets (from any season) on survey plots in the ParachutePiceance-Roan population of western Colorado, USA from 2009-2014. Values presented do not account for sample-unit level variation in pellet detectability.
PJ-Treatment Plotsa

PJ-Control (No Treatment) Plots

Sagebrush-Control Plots

1-Upper 2-Black
3-Ryan
MEAN
1-Dry
3-Stake
MEAN
1-Dry
2-Canyon
3-Black
MEAN
Galloway Sulphur
Gulch
± SE
Ryan
2-Eureka
Springs
± SE
Gulch
Creek
Cabin
± SE
b
b
b
c
Year
n = 49
n = 38
n = 48
N=3
n = 35
n = 50
n = 38
N=3
n = 54
n = 41
n = 42
N=3
d
2009
0.082
0.000
0.146
0.076
0.000
0.040
0.105
0.048
0.444
0.293
0.452
0.397
± 0.039 ± 0.000
± 0.051 ± 0.042
± 0.000
± 0.033
± 0.05
± 0.031
± 0.068
± 0.071
± 0.077 ± 0.052
2010
0.000
0.104
0.052
0.029
0.040
0.026
0.032
0.721
0.293
0.333
0.449
0.026
± 0.000 ± 0.026
± 0.044 ± 0.052
± 0.028
± 0.028
± 0.026
± 0.004
± 0.068
± 0.071
± 0.073 ± 0.136
2011
0.063
0.000
0.020
0.000
0.007
0.349
0.171
0.071
0.197
0.082
0.000
0.041
± 0.035 ± 0.041
± 0.000
± 0.020
± 0.000
± 0.007
± 0.073
± 0.059
± 0.040 ± 0.081
± 0.039 ± 0.000
2012
0.029
0.040
0.000
0.023
0.605
0.341
0.262
0.403
0.245
0.026
0.188
0.153
± 0.028
± 0.028
± 0.000
± 0.012
± 0.075
± 0.074
± 0.068 ± 0.104
± 0.061 ± 0.026
± 0.056 ± 0.065
2013
0.000
0.100
0.000
0.033
0.605
0.220
0.381
0.402
0.082
0.000
0.188
0.090
± 0.000
± 0.042
± 0.000
± 0.033
± 0.075
± 0.065
± 0.075 ± 0.112
± 0.039 ± 0.000
± 0.056 ± 0.054
2014
0.000
0.000
0.026
0.009
0.535
0.146
0.381
0.354
0.082
0.000
0.333
0.138
± 0.000
± 0.000
± 0.026
± 0.009
± 0.076
± 0.055
± 0.075 ± 0.113
± 0.039 ± 0.000
± 0.068 ± 0.100
2015
0.029
0.000
0.000
0.010
0.372
0.000
0.571
0.315
0.245
0.000
0.292
0.179
± 0.028
± 0.000
± 0.000
± 0.010
± 0.074
± 0.000
± 0.076 ± 0.167
± 0.061 ± 0.000
± 0.066 ± 0.090
a
Values in bold represent post-treatment data. Numbers before plot names refer to set (e.g., Set 1 = Upper Galloway, Dry Ryan, Dry Gulch, etc.).
b
n refers to no. of 30 x 30 m units sampled within the study plot.
c
N refers to no. of study plots contributing to the mean. Means in bold are for post-treatment plots, otherwise values represent pre-treatment data.
d
p = proportion of sample units per plots in which greater sage-grouse pellets were detected. Subscripts refer to year.

8

�Table 2. Preliminary data on observer variation in estimates of proportion of sample units with greater sage-grouse pellets (from any season) on survey plots from
repeat surveys in the Parachute-Piceance-Roan population of western Colorado, USA in 2013, 2014, and 2015. Values presented do not account for variation in
sample-unit level detectability of pellets. Surveys of each sample unit were conducted by a different observer within 20-30 minutes of each other on the same date.
PJ-Treatment Plots (post-treatment)
PJ-Control (No Treatment) Plots
Sagebrush-Control Plots
1-Upper
Galloway
n = 49a

2-Black
Sulphur
n = 38a

3-Ryan
Gulch
n = 48a

MEAN ±
SE
N = 3b

1-Dry
Ryan
n = 35

3-Stake
Springs
n = 38

MEAN ±
SE
N=3

0.000
± 0.000
0.000
± 0.000

0.013
± 0.013
0.033
± 0.033

0.465
± 0.076
0.605
± 0.075

0.195
± 0.062
0.220
± 0.065

0.238
± 0.066
0.381
± 0.075

0.299
± 0.084
0.402
± 0.112

0.026
± 0.026
0.026
± 0.026

0.009
± 0.009
0.052
± 0.024

0.535
± 0.076
0.488
± 0.076

0.146
± 0.055
0.098
± 0.046

0.381
± 0.075
0.548
± 0.077

0.354
± 0.113
0.378
± 0.141

0.286
0.000
0.271
0.186
0.057
0.000
0.000
0.019
0.465
± 0.065 ± 0.000 ± 0.064
± 0.093
± 0.039
± 0.000 ± 0.000
± 0.019
± 0.076
pBc
0.245
0.000
0.292
0.179
0.029
0.000
0.000
0.010
0.372
± 0.061 ± 0.000 ± 0.066
± 0.090
± 0.028
± 0.000 ± 0.000
± 0.010
± 0.074
a
n refers to the number of sample units per study plot.
b
N refers to the no. of study plots contributing to the mean.
c
p = proportion of sample units in which greater sage-grouse pellets were detected. Subscript refers observer (A or B).

0.024
± 0.024
0.000
± 0.000

0.548
± 0.077
0.571
± 0.076

0.346
± 0.162
0.315
± 0.167

2-Eureka
n = 50

1-Dry
Gulch
n = 43

2-Canyon
Creek
n = 41

3-Black
Cabin
n = 42

MEAN ±
SE
N=3

2013
pAc
pBc

0.224
± 0.060
0.082
± 0.039

0.000
± 0.000
0.000
± 0.000

0.167
± 0.054
0.188
± 0.056

0.130
± 0.067
0.090
± 0.054

0.000
± 0.000
0.000
± 0.000

0.040
± 0.028
0.100
± 0.042
2014

pAc
pBc

0.082
± 0.039
0.184
± 0.055

0.000
± 0.000
0.000
± 0.000

0.333
± 0.068
0.250
± 0.063

0.138
± 0.100
0.145
± 0.075

0.000
± 0.000
0.029
± 0.028

0.000
± 0.000
0.100
± 0.042
2015

pAc

9

�Table 3. Preliminary estimates of greater sage-grouse pellets per kilometer (from any season) encountered on transect plots in the Parachute-Piceance-Roan
population of western Colorado, USA from 2011-2014. Values presented do not account for variation in pellet detectability with distance.
PJ-Treatment Plotsa
PJ-Control (No Treatment) Plots
Sagebrush-Control Plots
Lower
Upper
Bar
Upper WagonMagnolia Cotton- Wagon- Lower Upper
Lower
Magnolia
WagonD
Magnolia Barnes
road
Bar D
South
wood
road
Bar D Bar D
Barnes
North
Sprague
road
Split
Control
Control Control Control
x2011b
0.10
0.00
0.00
0.00
0.00
0.00
0.00
18.82
13.51
6.46
5.61
0.00
0.00
13.78
c
X2011
0.03 ± 0.03
0.00 ± 0.00
11.10 ± 3.84
0.00 ± 0.00
x2012
X2012

2.86

x2013
X2013

0.31

x2014
X2014

2.14

x2015
X2015

0.00

kmd

9.80

0.27

0.00
0.75 ± 0.54
0.00
0.00
0.17 ± 0.11
0.00
0.00
2.24 ± 1.73
0.00
0.00
0.55 ± 0.53

0.00

0.63

25.51
-

2.02

0.00
0.31
0.58 ± 0.42

0.00

46.03

7.94
31.63
27.14 ± 7.97

22.96

0.52

0.00

2.89
-

0.00

0.00
0.00
0.00 ± 0.00

0.00

18.53

1.01
11.56
22.79 ± 12.93

60.03

0.10

8.96

3.91
-

0.00

0.00
0.10
0.03 ± 0.03

0.00

19.41

4.22
15.31
18.03 ± 5.98

33.16

0.10

2.67

0.00
-

0.00

0.00
0.10
0.03 ± 0.03

0.00

0.00

0.51
0.00
14.29 ± 14.12

56.63

7.52
7.64
9.58
6.36
5.88
6.44
12.52
9.80
4.68
6.80
5.92
5.88
5.88
(7.70)
(13.40)
a
Post-treatment data are in bold. Treatments on Lower Bar D and Upper Bar D occurred in Jan 2011. Data for Lower Bar D and Upper Bar D from 2011-2015
represent ~6, 18, 30, 42, and 54 mo post-treatment, respectively. Treatments on Lower Barnes were started by the Bureau of Land Management in summer
2007 and completed in summer 2008. Post-treatment data on Lower Barnes were collected opportunistically and are presented for comparison only.
b
x = no. pellet piles detected per km of transect.
c
X = mean no. pellet piles detected per km of transect across study plots within each treatment. Values for Lower Barnes are presented separately. The 2011
mean for Lower Bar D and Upper Bar D in 2011 represents post-treatment data and is presented separately from the 2011 mean for other PJ-Treatment plots
that had not yet been treated (Magnolia South, Cottonwood, Lower Wagonroad).
d
Total km of transect line surveyed per plot. Portions of transects on Cottonwood and Sprague unsuitable as greater sage-grouse habitat were eliminated in
2012, values in parentheses indicate km surveyed in 2011 only.

10

�Table 4. Timeline for research on greater sage-grouse response to pinyon-juniper removal in the ParachutePiceance-Roan population, western Colorado, 2008-2015.
Task
Identification of plots for pinyon-juniper removal
Remove pinyon-juniper on survey treatment plots (2009-2011)
Remove pinyon-juniper on transect treatment plots (2011)
Remove pinyon-juniper on new transect treatment plots (2014)
Pellet surveys (annually)
Prepare cumulative report (annually)
Prepare cumulative final report

11

Initiation
Summer 2008
Fall 2009
Fall 2011
Fall 2014
June-Aug
September
1 Sep 2015

Completion
COMPLETED
COMPLETED
COMPLETED
COMPLETED
COMPLETED
COMPLETED
COMPLETED

�Fig. 1. Map of study plot locations for the pinyon-juniper removal experiment and greater sage-grouse occupied range boundary (as
of 2012) in the northern portion of the Parachute-Piceance-Roan population of western Colorado, USA.

12

�Fig. 2. Trends in the proportion of sample units detected with greater sage-grouse pellets for individual survey plots (color-coded by
treatment) in the Parachute-Piceance-Roan population of western Colorado, USA, 2009-2015. Values presented do not account for
variation in sample-unit level detectability of pellets.

13

�Fig. 3. Trends in the mean proportion of sample units detected with greater sage-grouse pellets across survey plots (color-coded by
treatment) in the Parachute-Piceance-Roan population of western Colorado, USA, 2009-2015. Values presented do not account for
variation in sample-unit level detectability of pellets.

14

�Fig. 4. Trends in the no. of greater sage-grouse pellets counted per km of transect for individual transect plots (color-coded by treatment)
in the Parachute-Piceance-Roan population of western Colorado, USA, 2011-2015. Values presented do not account for variation in
detectability of pellets with distance from transect.

15

�Fig. 5. Trends in the mean no. of greater sage-grouse pellets counted per km of transect across transect plots within treatments (colorcoded by treatment) in the Parachute-Piceance-Roan population of western Colorado, USA, 2011-2015. Values presented do not account
for variation in detectability of pellets with distance from transect.

16

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                  <text>Colorado Division of Parks and Wildlife
September 2015-September 2016
WILDLIFE RESEARCH REPORT
State of:
Cost Center:
Work Package:
Task No.:

Colorado
3420
0660
N/A

Federal Aid
Project No.

N/A

:
:
:
:

Division of Parks and Wildlife
Avian Research
Greater Sage-grouse Conservation
Assessment of Greater Sage-grouse Response to
Pinyon-Juniper Removal in the ParachutePiceance-Roan Population of Northwestern
Colorado

Period Covered: September 1, 2015 – August 31, 2016
Author: B. L. Walker
Personnel: B. Holmes, B. Petch, T. Knowles, B. deVergie
All information in this report is preliminary and subject to further evaluation. Information MAY
NOT BE PUBLISHED OR QUOTED without permission of the author. Manipulation of these data
beyond that contained in this report is discouraged.
EXTENDED ABSTRACT
Greater sage-grouse (Centrocercus urophasianus) in the Parachute-Piceance-Roan (PPR) region of
western Colorado face at least two major potential stressors: projected habitat loss from energy
development and a long-term decline in habitat suitability associated with pinyon-juniper (PJ)
encroachment. Pinyon-juniper removal may be a useful mitigation tool to offset potential habitat losses
associated with energy development. Although pinyon-juniper removal is commonly used to improve
habitat for greater sage-grouse, no studies to date have quantified the timing or magnitude of how birds
respond to treatments. Since 2008, Colorado Parks and Wildlife (CPW) has cooperated with industry and
landowner partners to investigate the effectiveness of pinyon-juniper removal for restoring sage-grouse
habitat in the PPR. In fall 2008, I established nine “survey” study plots, arranged in three groups of three,
with each group consisting of a Sagebrush-Control plot, an untreated PJ-Control plot, and a PJ-treatment
plot. Treatments were completed on three survey plots in 2010 and 2011. Pellet surveys in summer from
2009-2015 indicated that the mean proportion of sample units containing pellets was consistently highest
on sagebrush control plots (range 0.197-0.449 across years), consistently lowest on plots with
encroaching pinyon-juniper (range 0.007-0.048), and increased on 2 of 3 treated survey plots (Ryan
Gulch and Upper Galloway) within 1-2 years after treatment. Fourteen transect plots were established in
fall 2010 and summer 2011, and two were established in summer 2014. Transect plots were surveyed for
pellets in summer though 2015. As expected, estimated mean pellet piles/km were low on the four PJControl plots for the duration of the study (range across years = 0.00-0.58 pellets/km) and on PJTreatment plots prior to treatment (mean = 0.03 pellet piles/km). Mean pellet piles/km were consistently
higher on all four Sagebrush-Control transect plots through 2014 (range across years = 11.10 - 27.14
pellet piles/km), but declined precipitously on 3 of 4 Sagebrush-Control plots in 2015 (Magnolia Control,
Upper Barnes Control, and Wagonroad Control). Pellet piles/km were also high on the Lower Barnes
transect plot 4-5 years post-treatment (13.78-25.71 piles/km), but declined 6-8 years post-treatment (0.003.91 pellet piles/km). Mean pellet piles/km has generally remained low on treated transect plots for four

1

�years after pinyon-juniper removal (range across years = 0.00 - 2.86 pellet piles/km) with the exception of
Upper Bar D in 2014 (8.96 pellet piles/km). Estimates of proportion of sample units with pellets (from
survey plots) and of pellet piles/km (from transect plots) also varied substantially among SagebrushControl plots within years and among years within plots, which suggests there is substantial variation in
pellets deposited within suitable habitat (i.e., over and above variation in counts due to observer bias). We
completed double-observer sampling on survey plots in 2013, 2014, and 2015 to estimate sample unitlevel detectability, and we completed distance sampling on transect plots in 2014 and 2015 to generate
distance-detection curves, and those analyses are in progress. We established and conducted pre- and
post-treatment surveys on two additional transect plots (Lower Galloway and Lower Ryan Gulch) in
summer 2014 and 2015. Analyses for this project are in progress. However, to date, sage-grouse response
to pinyon-juniper removal as measured by pellet surveys appears to be inconsistent, with pellet counts
increasing on only 2 of 8 plots within 4-5 years post-treatment. A recent experimental pilot project
suggests that sage-grouse may respond more strongly to both pinyon-juniper and serviceberry removal.
We are developing a proposal to expand the current project to include pinyon-juniper and/or serviceberry
treatments on existing and additional study plots starting in fall 2018.

2

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              <text>Greater sage-grouse (&lt;em&gt;Centrocercus urophasianus&lt;/em&gt;) in the Parachute-Piceance-Roan (PPR) region of western Colorado face at least two major potential stressors: projected habitat loss from energy development and a long-term decline in habitat suitability associated with pinyon-juniper (PJ) encroachment. PJ removal may be a useful mitigation tool to offset potential habitat losses associated with energy development. Although PJ removal is commonly used to improve habitat for greater sage-grouse, no studies to date have quantified the timing or magnitude of how birds respond to treatments. Since 2008, Colorado Parks and Wildlife (CPW) has cooperated with industry and landowner partners to investigate the effectiveness of PJ removal for restoring sage-grouse habitat in the PPR. In fall 2008, I established nine “survey” study plots, arranged in three groups of three, with each group consisting of a sagebrush control plot, an untreated PJ control plot, and a PJ treatment plot. Treatments were completed on the three treatment plots in 2010 and 2011.</text>
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              <text>Parachute-Piceance-Roan (PPR) region</text>
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              <text>Northwestern Colorado</text>
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              <text>Cost Center 3420 Avian Research. Work Package 0660 Greater Sage-grouse conservation&#13;
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