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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
Evaluating Lek-Based Monitoring and
Management Strategies for Greater Sage-Grouse in
the Parachute-Piceance-Roan Population in
Northwestern Colorado

Period Covered: September 1, 2013 – August 31, 2014
Author: B. L. Walker
Personnel: B. Holmes, S. Duckett, B. Petch, B. deVergie, J. T. Romatzke
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Information MAY NOT BE PUBLISHED OR QUOTED without permission of the author.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
Implementing effective monitoring and mitigation strategies is crucial for conserving populations of
sensitive wildlife species. Concern over the status of greater sage-grouse populations has increased rangewide and in Colorado due to population declines, range contraction, loss and degradation of sagebrush
habitat, and recent listing of the species as warranted but precluded under the Endangered Species Act.
Despite untested assumptions, lek counts are widely used as an index of abundance by state agencies to
monitor sage-grouse populations. Lek locations are also commonly used to identify and protect important
sage-grouse habitat. However, the use of lek counts and locations to monitor and manage sage-grouse
populations remains controversial because it is unknown how closely lek-count data track actual changes
in male abundance from year to year or if lek buffers are effective at protecting habitat for male sagegrouse during the breeding season. Colorado Parks and Wildlife is deploying solar-powered GPS satellite
transmitters on male greater sage-grouse to obtain data on male survival, lek attendance, inter-lek
movements, and diurnal and nocturnal habitat use around leks and conducting double-observer lek counts
to estimate detectability of males on leks during the breeding season in the Parachute-Piceance-Roan
population in northwestern Colorado. These data will allow us to evaluate whether current lek-based
monitoring methods provide reliable information about sage-grouse population trends and whether
current lek buffers are effective at protecting breeding males. Fourteen GPS males marked prior to 1 Sept
2013 were monitored for part or all of the 1 September 2013 - 31 August 2014 period. Field crews also
captured and deployed GPS transmitters on 21 additional males during the 2014 March-May breeding
season. One new potential lek was found by tracking GPS males in 2014, but at least 4 would have been
found had field crews not detected them earlier in the breeding season. Field crews conducted 93 standard
lek counts at 29 different leks, 28 unreconciled double-observer counts at 14 leks, and 46 paired ground
and helicopter counts at 21 leks in spring 2014. We obtained breeding-season location data for a total of
31 GPS males in spring 2014. We plan to capture additional GPS males on Chevron in March-April 2015.

1

�COLORADO PARKS AND WILDLIFE RESEARCH REPORT
EVALUATING LEK-BASED MONITORING AND MANAGEMENT STRATEGIES FOR
GREATER SAGE-GROUSE IN THE PARACHUTE-PICEANCE-ROAN POPULATION IN
NORTHWESTERN COLORADO
BRETT L. WALKER
PROJECT OBJECTIVES
(1)
(2)
(3)
(4)
(5)

Use locations of GPS males to find, verify, and count new leks
Estimate the number of known and unknown leks in the population
Estimate age-specific rates of male lek attendance
Estimate the frequency, timing, and distance of inter-lek movements by males
Estimate detectability of males attending leks using paired helicopter and ground counts and paired
ground counts
(6) Use these parameters in simulations to quantify how variation in lek attendance, inter-lek movements,
detectability, and count effort affect lek-count data and estimates of population trends collected using
standardized protocols
(7) Quantify male habitat use around leks to inform use of lek buffers
SEGMENT OBJECTIVES
(1) Capture and deploy solar GPS PTT transmitters on males during the spring 2014 breeding season to
reach a sample size of 25 GPS adults and 20 GPS yearlings.
(2) Use locations of GPS males to locate, verify, and count new leks in the study area in spring 2014
(3) Conduct standard counts, unreconciled double-observer counts, and paired ground and helicopter
counts at leks
(4) Enter and proof spring-summer field data
INTRODUCTION
Implementing effective monitoring and mitigation strategies is crucial for conserving populations
of sensitive wildlife species. Concern over the status of greater sage-grouse (Centrocercus urophasianus)
populations has increased both range-wide and in Colorado due to historical population declines, range
contraction, continued loss and degradation of sagebrush habitat, and recent federal listing of the species
as warranted but precluded under the Endangered Species Act in 2010 (Connelly et al. 2004, Schroeder et
al. 2004, CGSSC 2008, USFWS 2010). This concern is heightened in oil and gas fields due to recent
studies based on lek-count data that suggest negative impacts of development on sage-grouse abundance
(Holloran 2005, Walker et al. 2007, Doherty et al. 2010b, Harju et al. 2010, Tack 2010) and identification
of energy development as a threat factor in the eastern portion of the species’ range (USFWS 2010).
However, the use of lek-count and lek location data to monitor and manage sage-grouse populations
remains controversial. This uncertainty, in turn, has the potential to cause disagreement, controversy, and
conflict among agencies, industry, and other stakeholders where sage-grouse and oil and gas resources
overlap. For this reason, there is a crucial need to collect empirical data that evaluate whether current lekbased monitoring methods provide reliable information about population trends and whether current lekbased management strategies are effective for conserving greater sage-grouse in areas with expanding
energy development.
Each spring, male sage-grouse congregate on traditional mating grounds, or leks, to display and
mate with females (Schroeder et al. 1999). Males attending leks are then counted by observers on the

2

�ground or from aircraft following standardized protocols (Jenni and Hartzler 1978, Beck and Braun 1980,
Autenrieth et al. 1982, Connelly et al. 2000). Lek counts are the primary index used by all state wildlife
agencies in the western U.S., including the Colorado Division of Parks and Wildlife (CPW), to monitor
changes in sage-grouse abundance (Connelly et al. 2004, CGSSC 2008, WAFWA 2008, Fedy and
Aldridge 2011). Lek-count data are also commonly used to investigate how regional and range-wide
populations respond to changes in habitat and to anthropogenic stressors (e.g., Braun et al. 2002, Walker
et al. 2007, Aldridge et al. 2008, Doherty at el. 2010b, Harju et al. 2010, Tack 2010). However, the use of
lek-count data as an index of abundance has been called into question because the quantitative
relationship between lek counts and population size has never been clearly established. The use of lek
counts to measure population size rests on untested assumptions about that proportion of leks in the
population that are known and counted, the proportion of males that attend leks, the proportion of males
on leks that are detected by observers, and how often males move between leks during the breeding
season (Beck and Braun 1980; Applegate 2000; Walsh et al. 2004, 2010; CGSSC 2008; WAFWA 2008).
At present, too few quantitative data are available to estimate these parameters and their
associated variances. CPW initiated a project to estimate the number of leks and the proportion of known
leks in each population around the state using dual-frame sampling (Haines and Pollock 1998).
Preliminary data from the first three years of that project indicate that the proportion of known leks in
each population varies depending on their size and how well surveyed the population was prior to
sampling (P. Lukacs, CPW, unpublished data). Male lek attendance varies with age, time of day relative
to sunrise, date, weather, annual snowpack, renesting rates of females, predator activity, and human
disturbance, but previous studies of male lek attendance have not reported data in ways that allow us to
quantify how variation in male lek attendance influences annual lek-counts (Patterson 1952, Dalke et al.
1963, Rogers 1964, Hartzler 1972, Jenni and Hartzler 1978, Beck and Braun 1980, Autenrieth et al. 1982,
Emmons and Braun 1984, Ellis 1984, Dunn and Braun 1985, Connelly et al. 2000, Connelly et al. 2003,
Boyko et al. 2004, Walsh et al. 2004). Although standardization of lek-count protocols minimizes some
sources of variation in lek attendance (e.g., time of day, weather, date), a substantial amount of variation
is not accounted for. In the most rigorous studies to date, Walsh et al. (2004, 2010) emphasized the
importance of individual heterogeneity, age, sex, time of day, and date in determining lek attendance, but
because results were based on only one year of data from one small, geographically isolated population,
they concluded that additional research is needed before we can develop a comprehensive understanding
of annual and geographic variation in lek attendance. No published studies have quantified variation in
detectability of males on leks from year to year or how much detectability varies among observers, or
with weather, distance from lek, equipment used (binoculars vs. spotting scopes), or count method (e.g.,
ground vs. aerial counts) (Connelly et al. 2003, Walsh et al. 2004). Age-specific inter-lek movements by
males have been reported in several studies, with 4-50% of males known to have attended more than one
known lek during a single breeding season (Dalke et al. 1963, Gill 1965, Wallestad and Schladweiler
1974, Emmons and Braun 1984, Dunn and Braun 1985, Bradbury et al. 1989, Walsh et al. 2004), but the
effect of inter-lek movements on annual lek count data has not been quantified. Methodological
considerations may also affect lek counts. The sample of leks counted in any given year may depend on
which leks are accessible due to road conditions, snowpack, landowner permission, etc. This may bias
count data if access and attendance are correlated (e.g., if attendance is lower in areas with more roads).
Count effort (i.e., the number of counts per lek per breeding season) can also influence trend estimation
because often only the maximum count of males is recorded in state-wide databases. A maximum count
based on more visits is likely to be higher because additional visits are more likely to coincide with peak
male attendance (Walsh et al. 2004, CGSSC 2008, Fedy and Aldridge 2011).
Investigating the reliability of lek-count data for monitoring changes in actual population size is a
range-wide research priority for greater sage-grouse (Naugle and Walker 2007). Despite numerous
criticisms, lek-count data continue to be widely used as an index of abundance. Large decreases in lek
counts or disappearance of leks over large areas over time are thought to indicate population decline or

3

�range contraction in response to anthropogenic stressors (Walker et al. 2007, Aldridge et al. 2008,
Doherty et al. 2010b, Harju et al. 2010). The fact that state and regional core areas have been established
based largely on counts of males on leks and of lek density also reflects that state and federal agencies
consider higher lek counts, on average, to represent larger populations (CGSSC 2008, NRCS 2009,
Doherty et al. 2010a, Hagen 2010, State of Wyoming 2010). Indeed, lek counts remain the only widelyused method for monitoring populations of greater sage-grouse and there is general agreement among
wildlife professionals that they have inherent value for monitoring (Connelly et al. 2003, Naugle and
Walker 2007, CGSSC 2008). This raises an important question. Because state agencies and others
continue to use lek counts for monitoring, and because lek counts are affected to an unknown degree by
variation in attendance, inter-lek movements, and detectability, how much of a change in lek-count data is
required to reliably detect an actual change in male population size? In other words, how much do lekcount data bounce around due to unexplained variation in these parameters even when we follow
standardized count protocols even if male population size remained the same? Understanding the uses and
limitations of lek-count data will require both empirical field data from marked males and simulations
that illustrate how much standard lek counts vary when lek attendance, inter-lek movements, and
detectability are not accounted for.
Lek-based management strategies for greater sage-grouse also require evaluation. Leks are
typically centrally located within nesting areas and often overlap with other seasonal habitats (Connelly et
al. 2000, Doherty et al. 2010c, Aldridge et al. 2011, Fedy et al. 2012), so lek locations are commonly used
to help identify and protect important sage-grouse habitat. There is also concern that disturbance at leks
may cause abandonment of those leks or reduce rates of nest initiation or reproductive success (Lyon and
Anderson 2003, Holloran et al. 2010). For these reasons, state and federal agencies typically recommend
restrictions on surface occupancy, timing restrictions during the breeding season, or both, within a certain
buffer distance around leks in oil and gas fields (CGSSC 2008). Agencies have also delineated “core
areas” or “priority breeding habitats” by placing buffers around leks that meet minimum male count and
lek density criteria (e.g., CGSSC 2008, Doherty et al. 2010a, Hagen 2010, State of Wyoming 2010).
However, these types of lek-based management strategies are subject to two major criticisms.
First, lek-based management strategies incorrectly assume that all lek locations are known
(CGSSC 2008). New lek locations are discovered each year in northwestern Colorado (CPW, unpublished
data) and hundreds of new leks have been discovered in the past decade throughout the species’ range
(WAFWA 2008). Because lek-based oil and gas lease stipulations (e.g., lek buffers) can only be applied
to leks whose locations are known, the presence of undiscovered leks in oil and gas fields may result in
inadequate protection for populations. Although lek-based monitoring data can be adjusted to account for
unknown leks, lek-based management strategies cannot. For this reason, appropriately managing sagegrouse in oil and gas fields using a lek-based approach requires estimating the total number of leks in the
field, estimating the proportion of those leks that are known, identifying where unknown leks are most
likely to occur, and finding, verifying, and counting new leks.
A second criticism of lek-based management strategies is that they have not been empirically
validated for specific local populations. Current federal oil and gas leases typically contain stipulations
for either no surface occupancy (NSO) or restricted surface occupancy (RSO) within certain buffer
distances around leks. Historically, the Bureau of Land Management implemented a 0.25-mi. NSO or
RSO buffer around leks to minimize disturbance to lekking males and to protect habitat that males use
during the breeding season, with the overall intention of minimizing long-term population declines and
preventing extirpation in areas with development (CGSSC 2008). However, a 0.25-mi. stipulation has no
credible scientific basis (p. B-5, Appendix B, CGSSC 2008). More recently, a review of six range-wide
studies of male diurnal habitat use and movements during the lekking season suggested that a 0.6-mile
buffer around leks may be more appropriate (p. B-6, Appendix B, CGSSC 2008). This criterion is now
recommended by state and federal agencies in Colorado (CGSSC 2008). However, some studies have

4

�questioned whether lek buffers designed to protect males are adequate to prevent sage-grouse populations
in oil and gas fields from declining below desired thresholds (Holloran 2005, Walker et al. 2007, Harju et
al. 2010). In contrast, other authors have questioned whether it is appropriate to apply a one-size-fits-all
lek buffer based on range-wide data to local populations with different topography and vegetation and
subject to different types and intensities of energy development (Harju et al. 2010). Lek buffers are also
sometimes criticized for including habitat within the buffer that is clearly unsuitable for sage-grouse or
being so large so as to effectively preclude energy development. No studies to date have empirically
tested how large buffers actually need to be to protect habitat for males during the lekking season or
quantified what level of protection buffers of different sizes provide for year-round habitat in any given
local population. It remains unclear whether a 0.6 mi. buffer is too large, adequate, or too small, or
whether other buffer sizes would be more appropriate. For this reason, field research is needed that
quantifies the size of lek buffer required to adequately protect male sage-grouse during the breeding
season in local populations subject to oil and gas development by intensively tracking both diurnal and
nocturnal habitat use of males around leks. These data will allow us to develop scientifically defensible
recommendations regarding the appropriate size and use of lek buffers for specific oil and gas fields.
Recent technological advances have led to production of 30 g, solar-powered, global positioning
system (GPS) satellite transmitters suitable for use with greater sage-grouse. GPS transmitters have
several advantages over traditional VHF transmitters that make it possible to generate the data needed to
resolve lek-based monitoring and management issues. GPS transmitters can be programmed to record
multiple locations at specific times of day, logistical problems that prevent crews from locating birds on
the ground (e.g., bad weather, poor road conditions, truck breakdowns, lack of access, etc.) do not bias
data collection, data are gathered without disturbing the bird or its flock mates, and the units provide the
high-resolution data needed to estimate male lek attendance, inter-lek movements, and diurnal and
nocturnal habitat use around leks.
I propose a three-year research project to evaluate greater sage-grouse lek monitoring and
management strategies in oil and gas fields in the Parachute-Piceance-Roan population of northwestern
Colorado using GPS transmitters deployed on males. This study will directly complement a second,
overlapping research project sponsored by CPW and Colorado State University (Fort Collins) and funded
by Exxon-Mobil to examine alternative methods of population monitoring, including: dual-frame
sampling of leks and non-invasive genetic sampling for sex ratio and population estimation.
Data collected on this project will allow us to judge the reliability of lek-count data for producing
defensible estimates of male population size and trend over time as energy development proceeds, thereby
directly informing whether sage-grouse management and conservation efforts by industry, agencies, and
other stakeholders in oil and gas fields are effective. This research will also provide insight into the
ecological and methodological factors that need to be considered when collecting and analyzing lek-count
data and appropriate uses and limitations of lek-count data for monitoring. This research will also provide
local data that landowners and managers can use to test and make informed decisions about the
effectiveness of lek buffers for mitigation. These expected results, in conjunction with studies in other
parts of NW Colorado, will in turn have both state-wide and region-wide implications for monitoring and
managing both greater sage-grouse and Gunnison sage-grouse (C. minimus).
STUDY AREA
The study area encompasses the current occupied range of greater sage-grouse in the ParachutePiceance-Roan (PPR) population (Fig. 1), one of 7 geographically distinct populations in northwestern
Colorado. The PPR population is experiencing rapid changes to sage-grouse habitat due to expanding oil
and gas development (CGSSC 2008). Ownership within occupied range in the PPR is 65% private
(primarily ranches and energy companies), 33% federal (primarily Bureau of Land Management), and 2%

5

�State of Colorado (PPR-GSGWG 2008). Based on 2007 lek-count data, the PPR population represents
approximately 4% of the total population of greater sage-grouse in Colorado (CGSSC 2008). The
population appears to have experienced a recent decline since 2006; counts in 2011 totaled 106 males on
32 leks, for an average of 3.3 males per active lek (CPW, unpublished data; Neubaum 2011). Count data
in 2011 are approximately one-half the known high count of 204 males on 28 leks in 1976 (7.3
males/active lek; Krager 1977) or the high count of 226 males in 2006 (PPR-GSGWG 2008). Many of the
88 known lek locations in the PPR population are unoccupied (Fig. 1), but new leks have been found
recently, in part due to greater aerial survey effort since 2005 and intensive tracking of marked birds by
research crews since 2006 (PPR-GSGWG 2008, Apa 2010).
Sage-grouse in the PPR population inhabit the tops of ridges and plateaus dominated by mountain
big sagebrush (Artemisia tridentata vaseyana) and mixed sagebrush and “mountain shrubs” (e.g.,
serviceberry, Amelanchier spp.; Gambel oak, Quercus gambelii; snowberry, Symphoricarpus sp.;
antelope bitterbrush, Purshia tridentata; and mountain mahogany, Cercocarpus sp.). These habitats occur
between 6,500-9,000 feet and are often interspersed with patches of aspen (Populus tremuloides) and
conifer forest. These ridges and plateaus are intersected by relatively steep drainages and bordered at
lower elevations by pinyon-juniper woodland. Birds only use certain areas that have suitable local
topography and local- and landscape-scale vegetation features (Apa 2010, Walker 2010), so sage-grouse
habitat in the PPR is actually more restricted than indicated by the occupied range boundary (CGSSC
2008). Due to the elevation, males display slightly later in the PPR than in lower-elevation populations.
Males in the PPR typically begin to display on leks in late March or early April (depending on the year)
and may continue to display though early June, with a peak of lek activity from mid-April to mid-May.
Snowpack depth and duration in the PPR varies from year to year, and strutting is thought to start later
following winters when snowpack lasts longer, presumably because snow covers potential nest sites and
delays forb growth important for females prior to incubation (Gregg et al. 2008).
METHODS
Capture and Handling
I plan to capture and attach 30-g, rump-mounted solar-powered GPS PTT satellite transmitters
(Northstar Science and Technology, King George, VA) on adult male greater sage-grouse in OctoberNovember and on yearling males in March of each year. Adult males will be captured in fall prior to the
onset of the breeding season to prevent biasing data on lek attendance the following spring (Walsh et al.
2004). Yearling males will be captured in March-April to allow them to reach larger body size prior to
deploying the transmitter, thereby reducing the chance that they will outgrow the harness. Trapping effort
will be distributed across the population so that it is, as much as possible, proportional to, and
representative of, the amount of local breeding habitat present as identified in seasonal habitat models
(Walker 2010). I selected 30-g transmitters because they have larger battery capacity than 22-g models,
which decreases risk of transmitter failure under low-light conditions (e.g., during winter, or if birds
burrow under the snow during storms). A 30-g GPS transmitter with harness (38 g total) represents ~1.11.9% of male body mass, depending on age. GPS males will also receive individually numbered
aluminum leg bands (size 16). Transmitters from birds that die may be recovered, cleaned, refurbished,
and redeployed as necessary to maintain sample sizes.
Capture and handling methods will follow standard CPW operating procedures established for
sage-grouse, with the exception that field crews will deploy GPS transmitters (as described in detail
below), and the decision whether injured birds will either be released or euthanized will be made by the
PI rather than transporting birds back to Fort Collins. No known rehabilitators in western Colorado
currently have the facilities to care for wild sage-grouse. Crews will capture males using net launchers
(Giesen et al. 1982), night-time spotlighting and hoop-netting (Wakkinen et al. 1992), walk-in traps
modified for sage-grouse (Schroeder and Braun 1991), Super Talon® net guns (Advanced Weapons

6

�Technology, La Quinta, CA), MagNet® net guns (Wildlife Capture Services, Flagstaff, AZ), or throw
nets, all of which have been approved for capture of greater sage-grouse in this population by CPW’s
Animal Care and Use Committee.
Sample Size
I selected a sample size of 20 adults and 15 yearlings for the first year of the study as a
compromise between getting sufficient data to ensure I can estimate the parameters needed and
minimizing impacts on the population should GPS transmitters have any possible detrimental effect on
males. Once data are available from a concurrent study regarding impacts of GPS transmitters on male
survival and for variance estimates for lek attendance, inter-lek movements, and detectability parameters,
I will conduct a power analysis to assess whether sample sizes should be increased in the future.
Deploying more than 35 GPS transmitters in the first year might be considered an unacceptable risk to the
population and may not be supported by stakeholders or cooperators until we know from other studies
whether GPS transmitters affect annual survival. I will increase sample sizes in the second and third years
of the project to 30 adults and 25 yearlings if the first year of data from PPR or if data from an existing
study in the Hiawatha region in northwestern Colorado indicate no obvious impacts of GPS transmitters
on male survival. I will report survival rates of GPS males in my annual reports to the ACUC committee
as those data become available. I will also report the number of injured or euthanized birds to ACUC.
This will help establish standardized protocols for the use of GPS transmitters with greater sage-grouse.
Band-recapture data suggest that survival rates of male sage-grouse vary annually and by age (0.635 ±
0.034 SE for yearlings vs. 0.368 ± 0.007 SE for adults; Zablan et al. 2003). I plan to deploy 20
transmitters on adults in fall because I anticipate some level of transmitter failure and mortality (possibly
as many as 5 units) prior to the spring breeding season, thereby resulting in approximately equal sample
sizes of adults and yearlings (~15 each) in spring 2012.
GPS Transmitter Attachment
I will use a rump-mounted, leg-loop harness attachment for GPS transmitters based on the
method B design described in Bedrosian and Craighead (2007) modified for sage-grouse (Figs 2, 3). GPS
transmitters cannot be used with necklace collars because solar cells under the neck receive insufficient
sunlight to charge the battery. A thin layer of neoprene (1/8”) is glued to the bottom side of the transmitter
to ensure that contact between the transmitter material and the bird’s lower back is padded and insulated.
The harness material is 0.55-cm (0.25-inch) wide, brown Teflon ribbon (Bally Ribbon Mills, Bally, PA).
A 12-16 cm length of 0.55-cm wide elastic cord is sewn into the center of a 75-cm (36-inch) length of
Teflon ribbon such that 4-6 cm of expandable Teflon ribbon extends out from the attachment points on
either side (Fig. 2). The elastic gives the harness flexibility when the bird extends its legs during take-off
and when males are displaying. Yearling harnesses were sewn with more elastic (16 cm) to accommodate
any possible increase in body size over time. Transmitters are manufactured with a medium-brown, sandtextured finish to reduce reflected light and the sides, front, and back are painted with a camouflage
pattern to decrease visibility to predators (Fig. 3). Problems with back feathers near the front and sides of
the transmitter covering the solar panel and leading to transmitter failure led us to develop an improved
design that uses thicker neoprene (1/4”) and longer and wider neoprene under the front half of the
transmitter (Fig. 3). The new design should prevent feathers in front of, and on the sides of, the
transmitter from covering the solar panel. I selected a light, flexible antenna to minimize interference with
the bird’s upright tail display while strutting. The transmitter is mounted on the bird’s lower back
centered between the legs (as seen from behind and as seen from the side of the bird) with the antenna
extending toward the rear above the tail (Fig. 4). Harness leg-loops on each side are fitted while the bird
is held on its side to allow careful placement of the Teflon ribbon under body feathers and under the leg.
The leg-loop is fitted down, around, and underneath the legs, then back up and through the attachment
loops on the transmitter, then secured in place using a small section of 0.55-cm (0.25-inch) diameter
copper tubing as a crimp (Fig. 2). The final fit is checked with the bird held in a normal, standing
position. Transmitters are fitted as loosely as possible, but just snugly enough to prevent birds from

7

�dropping transmitters. Copper crimps quickly become tarnished with exposure to the elements, but as a
precaution, crimps are marked with black ink before release to reduce reflection. Field crews will trim the
Teflon ribbon at an angle and leave just enough excess ribbon on each side (~3 cm) to allow refitting or
enlargement of the harness should it become necessary. The end of the excess ribbon is dabbed with
Superglue® (Super Glue Corporation, Rancho Cucamonga, CA) to prevent fraying. The life span of the
exposed Teflon ribbon and elastic inserts has not been experimentally tested, but harnesses from
transmitters on females recaptured after &gt;30 months in the field show no signs of fraying, wear, or
deterioration.
The GPS transmitters are solar-powered and may last for 3-5 years, which is longer than the life
span of almost all male sage-grouse (Zablan et al. 2003). All transmitters are pre-programmed to collect 8
locations per day from March-May to gather data on early morning lek attendance (6 am, 6:30 am, 7 am,
7:30 am, 8 am), mid-day feeding/loafing areas (12 pm), evening feeding areas (6 pm), and night roost
locations (12 am). Transmitters are programmed to collect two locations per day at 12 pm and 12 am from
June-Feb to capture basic patterns of seasonal habitat use and movements while reducing demand on the
battery during low-light conditions encountered in fall and winter. I anticipate studying males for multiple
seasons, so I do not plan to remove GPS transmitters from males until summer 2014 following the final
breeding season. I will clean and redeploy GPS transmitters recovered from mortalities on additional
males as needed to maintain sample sizes. I was trained in attachment techniques in the field by Bryan
Bedrosian, who has used GPS transmitters with raptors, corvids, and sage-grouse (Craighead and
Bedrosian 2009). I have successfully deployed GPS transmitters on 34 females and 35 males from March
2009 - March 2011.
There is some concern that GPS transmitters attached using a rump-mount harness may
negatively affect male survival, movement, or strutting. Annual survival of female greater sage-grouse
with GPS transmitters in northwestern Colorado was lower than females with VHF transmitters from 5
April 2009 to 23 June 2010 (0.494 ± 0.109 SE, n = 40 for VHF vs. 0.346 ± 0.109 SE, n = 52 for GPS) (B.
Walker, unpublished data). However, harnesses have since been made more flexible by sewing in extra
elastic, transmitter camouflage was improved (Fig. 3), and 30-g transmitters (38 g including harness and
crimps) represent proportionally less of the body mass of males (1.1-1.9%, depending on male age;
~2000-2400 g for yearling males, ~2800-3300 g for adult males; Beck and Braun 1978) than of females,
all of which should minimize possible negative impacts of transmitters. CPW deployed GPS transmitters
and color-bands on male greater sage-grouse in fall 2010 and spring 2011 as part of a similar study in the
Hiawatha Regional Energy Development project area in northwestern Colorado. Preliminary data from
the Hiawatha project indicate that a lower proportion of color-banded GPS males (0.59 of 32 adults; 0.44
of 18 yearlings) were resighted at least once on leks in spring 2011 than were color-banded only males
(0.77 of 43 adults; 0.60 of 5 yearlings), but estimates of annual survival of color-banded GPS males
versus color-banded only males won’t be available until after resighting data are collected in spring 2012.
A recent comparison of movement data from greater sage-grouse females with GPS vs. VHF transmitters
throughout Wyoming found no evidence of a difference in distance traveled during seasonal or migratory
movements (Fedy et al. 2012). Observers at leks in Hiawatha in spring 2011 noted that transmitter
antennas are pushed to the side during strutting and do not appear to interfere with the bird’s upright tail
display. Moreover, a paired comparison of strutting behavior on leks detected no difference in mean
display rate between males with GPS transmitters (4.06 displays/min ± 0.45 SE) and adjacent, unmarked
males in the same age class (3.61 displays/min ± 0.36 SE) (paired t-test; t = 0.254, n = 43).
Several males with GPS transmitters have disappeared both in this study and in another study in
the Hiawatha region. To improve our ability to determine the fate of GPS males whose transmitters stop
transmitting and to relocate transmitters that disappear, we added a 5.3-g auxiliary VHF mortality microtransmitter (Advanced Telemetry Systems, Model A2720, Isanti, MN) on the underside of most GPS
transmitters starting in fall 2012 (Fig. 3).

8

�Lek and Lek Attendance Definitions
I define a lek as any restricted geographic area within which ≥ 2 males have displayed during the
breeding season in ≥ 2 years (over any number of years), which is consistent with previous state-wide and
range-wide definitions (Connelly et al. 2000, 2004; CGSSC 2008). I use this definition to ensure that
small leks and “satellite” leks are included, but that locations where males do not consistently display are
excluded (i.e., one-time use locations). It is possible that one-time use locations are important in small
populations like the PPR (S. Duckett, CPW, pers. comm.). If so, this will cause us to underestimate lek
attendance because we are unable to confirm strutting at these locations based on a single morning GPS
location. The status of a lek may be active or inactive in any given year. Leks used by displaying males at
least once within the past 5 years are considered active (CGSSC 2008). Newly-discovered leks &gt; 500 m
from all other known leks will be designated as potential leks. If those locations are documented to have
displaying males in ≥ 2 years, they will be classified as new leks and assigned a name based on local
geography. Field crews will delineate a “count boundary” for each known lek. The count boundary
represents the specific perimeter within which males would be visible and available for counting by
observers during any given count. The purpose of establishing a count boundary is to ensure that the
geographic area of observation for each lek is consistent over time. This prevents the characteristics of
specific leks (e.g., their size, location, topography, etc.) from changing over time. This count boundary
will necessarily be larger than the outer perimeter around displaying males on any given date because: (a)
observers can typically see and count males over an area larger than just the area where displaying males
are found, (b) males may shift the location where they strut slightly from day to day, and (c) observers
typically adjust the location from which they count males from day to day to maximize their ability to
obtain complete counts of males.
It is also important to unambiguously define lek “attendance” because some males use habitat
near leks, but they may or may not be within the area that can be counted by observers. I define lek
attendance as the presence of the male at a location that falls inside within 26 m the count boundary (i.e.,
visible and available for counting by observers) at any time during the standard count period (0.5 hrs
before sunrise to 1.0 hr after sunrise) during the breeding season (15 March – 15 June). A male is not
considered to have attended a lek if the male is either: (a) outside the count boundary (i.e., not visible and
unavailable for counting) during the standard count period, or (b) inside the count boundary at a time
other than during the standard count period or outside the breeding season. Lek attendance of GPS males
should be straightforward to assess when resighters are present, but there may be some ambiguity about
lek attendance for GPS males that are not directly observed (those that attend leks at which no observers
are present). The positional accuracy of locations derived from GPS transmitters is typically ≤ 26 m. GPS
males with early morning locations within 26 m of the count boundary will be considered inside the count
boundary.
Lek Counts
CPW lek-count protocol instructs observers to obtain a maximum count of males by conducting
repeated counts 5-10 minutes apart over a 30-minute period between 0.5 hr before and 1.0 hr after sunrise.
Although no specific guidelines are given for the distance at which leks should be surveyed, an informal
survey of NW region biologists and wildlife managers suggest that they typically count leks from 50-400
m, depending on topography, access, and to avoid disturbing birds at the lek. Observers use whichever
optics are required to obtain a reliable count (binoculars or spotting scope) and whichever mode of
transportation (truck, ATV, on foot) gets them close enough to the lek to count it.
Field crews will focus on collecting double-observer count data at leks at least once a week. In
the PPR, not all leks are accessible early in spring due to snowpack and road conditions. Observers will
work in pairs, and each pair will conduct a 30-minute lek count during the standard count period at two
leks per day, if possible. Each 30-minute visit to a lek will be divided into six 5-minute scan intervals.

9

�Counters will follow CPW count protocols and record the maximum number of yearling and adult males
and females counted during each 5-minute interval. The goal of each counter is to get an accurate count of
yearling and adult males and females during each scan interval and to record any GPS males present on
the lek. Counters will also record when and how many birds of each sex arrive or leave the lek during
each interval. Counters will alternate between using a spotting scope and binoculars during each scan
interval. Each observer will be allowed to scan the lek multiple times within each 5-minute interval to be
consistent with how surveys are conducted by CPW biologists and wildlife managers. All observers will
be trained in standard lek-count protocols and aging and sexing; they will practice counting under
supervision prior to collecting field data; and they will collect data on standardized forms to ensure
consistency. All counts will be conducted from within a realistic distance from leks, depending on
topography and optics (50-400 m), and all counters will record the distance (m) to the approximate lek
center using a laser rangefinder. All observers will conduct counts using the same standard make and
model of 10x binoculars and 20-60x zoom spotting scopes.
Aerial lek counts will be conducted from a helicopter (the pilot plus an observer). Aerial
observers will focus first on counting birds at leks simultaneously being counted by ground crews, and
then conduct standard counts and search for potential new leks at specific locations indicated by GPS
male location data. Aerial observers will record the total number of males at a lek but cannot distinguish
adults from yearlings.
Objectives
Objective 1: Use locations of GPS males to find, verify, and count new leks – Early morning
locations of GPS males will be compared against locations of known leks as the data come in to identify
potential lek locations. Males that make ≥ 2 early morning visits to the same approximate location (within
100 m) on consecutive mornings during the breeding season will be considered to have visited a potential
lek location. The field crew will then visit those locations or they will be checked from the air at least
once during the next 7 days under suitable weather conditions to document whether displaying males or
their sign (e.g., pellets, tracks, feathers) are present. If displaying males or sign are present at a newly
discovered lek, then the new lek will be added to the list of regularly counted leks following standard
protocols, and the count boundary determined as soon as practicable. A GPS male that previously used a
location within the count boundary of a newly discovered lek during the standard morning count period
will be considered to have attended that lek on that date.
Objective 2: Estimate the no. of known and unknown leks in the study area – Data from GPS
males will be used in a mark-recapture framework to estimate the number of leks in the study area. Visits
by marked GPS males can be used to “capture” leks and subsequent visits by marked birds to that lek
constitute “recaptures” of that lek. Leks not attended by GPS-marked males are considered un-marked
leks. Recapture histories for individual leks can then be derived and analyzed using an appropriate markresight model (Bartmann et al. 1987, Bowden and Kufeld 1995) in program MARK (McClintock et al.
2008, White 2010).
Objective 3: Estimate age-specific lek attendance by males – I will use the GPS male dataset to
estimate lek attendance as a function of male age and variables that can be measured without directly
observing attending males. I will compare early morning locations of males with GPS transmitters against
count boundaries for all known lek locations to determine whether or not GPS males attended leks (see
definition of “attending a lek”, above). I can then estimate daily rates of lek attendance for each male
using logistic regression. Field crews will document all major weather events that could influence male
attendance throughout the field season (e.g., storms, high winds, etc.). Daily lek attendance for males will
be modeled as a function of date, weather, previous day weather, nearest lek size, and the male’s previous
lek attendance (as a measure of a male’s prior reproductive effort). Overall lek attendance for each
individual over the season will be calculated by summing the total number of days that each bird attended

10

�a lek and dividing by the total number of days for which each individual was alive and its early morning
location was known. Males that move outside the study area prior to the breeding season will be excluded
from the analysis.
Objective 4: Estimate rates of inter-lek movements of males – I will use location data from GPS
males overlaid with locations of count boundaries for all known active leks to estimate the frequency,
timing, and distance of inter-lek movements by yearling and adult males.
Objective 5: Estimating detectability of males on leks – I will use an unreconciled, independent,
double-observer approach to estimate detectability from lek-count data (Riddle et al. 2010). Standard
double-observer and removal models require that observers match or reconcile specific individual animals
that were or were not detected by each observer (Nichols et al. 2000). However, this is typically
impossible to do with lek counts because there may be numerous males on the lek and most males are
unmarked and cannot be individually identified. Unreconciled double-observer models use raw maximum
counts of the number of individuals detected (in each age class or from both age classes combined) from
each of two independent observers to generate a site history for each observer on each count (e.g., 13, 15)
(Riddle et al. 2010). Site histories are then analyzed using the repeated-counts hierarchical model of
Royle (2004) in program PRESENCE, with the only difference that data from each observer are
considered independent “visits” (Riddle et al. 2010). Benefits of this approach are that leks do not actually
have to be visited twice on separate occasions, and the closed population assumption is met because
surveys are conducted at the same time (Riddle et al. 2010). The method may require using a negative
binomial or zero-inflated Poisson distribution in place of a Poisson distribution if data are overdispersed
(Riddle et al. 2010). This estimator may become unstable if detectability is low (P. Lukacs, CPW, pers.
comm.). However, I anticipate relatively high detectability because observers typically position
themselves to maximize their ability to detect males attending the lek.
The counting protocol outlined above (under Lek counts) results in dataset with six repeated
counts from the same lek on each date for each counter for each age class of males and for females, with
three of the six counts by each counter done with a spotting scope and three with binoculars. Counters
will record distance to approximate lek center and presence or absence of snow cover on the count as well
as predator activity, weather (temperature, wind speed, precipitation, visibility, illumination), and any
other disturbances (e.g., vehicle traffic, planes, deer/elk, etc.) at the end of each 5-minute interval.
Predator activity will be broken into three classes (no predator detected; predator visible from the lek;
predator on, over, or attacking males) based on observations of potential predators of adult sage-grouse
(eagles, hawks, falcons, owls, coyote, red fox, bobcat, mountain lion, etc.) near the lek. Covariates in the
analysis of site histories will include a random effect of lek, and fixed effects of lek size (i.e., max no. of
males counted), distance from lek, optics used (binoculars vs. spotting scope), observer, predator activity,
weather, and an interaction between optics and distance from the lek. Because the data consist of repeated
counts from the same lek within and among days, this dependence will have to be addressed using a
repeated-measures approach.
To estimate the effect of counting males from the air on detectability, I will compare maximum
counts of males conducted on the same date at the same time from a single observer on the ground against
a single observer in a helicopter using an unreconciled double-observer approach. Crews will attempt to
conduct at least 3 paired counts per lek per year during the standard lek-count period in April, at least for
leks that are accessible to ground-based observers. Ground observers will count leks over the full standard
morning count period (0.5 hrs before sunrise to 1.0 hrs. after sunrise) and will note when the helicopter
flew over and any changes in the count (e.g., departures) or behavior (e.g., crouching) of males caused by
the helicopter that might affect count data or male detectability. Observers will continue to count until the
end of the standard morning count period (1.0 hr after sunrise) to assess how helicopter flyovers would
affect count data collected from the ground following a flyover (McRoberts et al. 2011).

11

�I will first compare the maximum count from the helicopter versus the maximum count from
either observer on the ground collected over the 30-minute count period prior to arrival of the helicopter.
Helicopters tend to flush males off leks and are likely influence counts made immediately after the
flyover (S. Duckett, CPW, unpublished data; McRoberts et al. 2011), so I will only include data collected
prior the helicopter’s arrival in the comparison. In this case, the difference in detection probability among
observers (ground vs. air) represents the difference in detectability on a typical standard ground count
versus a typical count from a helicopter. This comparison is appropriate because ground counts based on
data from a 30-minute count period and flight counts based on data from a 1-3 minute count period are
recorded with equal weight in statewide count databases.
I will also compare counts recorded from the helicopter versus just the last count recorded on the
ground in the 3 minutes prior to arrival of the helicopter. This allows us to estimate the effect of just the
count method (helicopter vs. ground) based on a similar time period.
It is unclear whether detectability will be higher or lower on helicopter counts. Detectability may
be lower because data are derived from only 1-2 passes over 1-3 minutes rather than a full 30-minute
period. However, it is possible that observers in helicopters may count more males if topography prevents
observers on the ground from detecting hard-to-see males around the periphery of a lek.
Objective 6: Simulate lek-count data – I will use estimates of the proportion of known vs.
unknown leks in the population, age-specific means and variation in male lek attendance, the frequency
and distance of inter-lek movements, estimated detectability, and variation in lek-count effort in
conjunction with important covariates (e.g., time of day, date, weather, etc.) to simulate how much lek
counts are likely to vary, even in the absence of population change, when conducted according to
standardized protocols. I will simulate data for the estimated total number of leks in the population. The
number of males in the simulated population will be set at a value equal to the maximum count of
yearling or adult males at each lek during the period of peak attendance for each age group divided by
age-specific detectability during that period. I can use these data to simulate what proportion of the
simulated population of adult and yearling males would actually attend each lek during each time period
of the morning on each day of the breeding season each year. I would then run scenarios using this
simulated dataset with realistic combinations of measured and unmeasured variables that influence
detectability (e.g., time of day, optics, distance from lek, weather, and number of counts per season).
Scenarios would include counts conducted: (a) under more restrictive (0.5 hrs before to 0.5 hrs after
sunrise) or less restrictive (0.5 hrs before to 1.0 hrs after sunrise) time of day requirements; (b) with
binoculars versus spotting scope; (c) at various distances from leks; (d) in good versus marginal weather
conditions; (e) with a variable number of counts per season (from 1 to 6) on randomly selected dates at
least a week apart (to mimic data contained in state databases); (f) with varying proportions of leks
counted or a biased sample of leks counted to mimic normal problems with access encountered in the
field. Simulations will be set up in program R (version 2.11.0, R Development Core Team 2010).
Objective 7: Test 0.6-mile lek buffer – During the breeding season, I will measure distances of
three off-lek locations per day (at noon, 6 pm, and midnight) for each GPS male to the center of the lek
attended that morning, the lek most recently attended, the nearest active lek (as recorded in CPW
databases or by field crews), and the lek attended on the next visit. During other seasons, I will measure
distances of two off-lek locations per day (at noon and midnight) for each GPS male to the center of the
nearest known active lek (as recorded in CPW databases or by field crews). I will then calculate the
proportion of off-lek locations (for each portion of the day) that fall within specific distances of dissolved
buffers around the centers of known active leks to test the effectiveness of the current 0.6-mi. NSO/RSO
stipulation for lek buffers and to make recommendations on the most efficient buffer size to use to protect
specific proportions of the population. It may also be possible to use a kernel or bivariate normal mixture

12

�model to estimate the probability of males using areas within a specific distance around leks (D. Walsh,
CPW, pers. comm.). I will also compare the effectiveness of conserving areas within different circular lek
buffers to similar areas of high priority breeding habitat of similar size already identified using VHF
locations of females (Walker 2010).
RESULTS AND DISCUSSION
Captures
Field crews captured and deployed GPS transmitters on 21 males during the March through May
2014 breeding season using hoop nets, MagNet® net guns, and lightweight throw nets.
GPS Transmitters
Of the 65 males successfully deployed with GPS transmitters from March 2012 through May
2014, 23 were still alive with functioning transmitters, 3 slipped out of their transmitters, 25 are known to
have died, and the status of 14 could not be determined as of 31 Aug 2014. Feathers covering the solar
panel were a significant factor in transmitter failure with our original transmitter design (Fig. 2). In fall
2012, we developed and implemented an improved transmitter design using thicker neoprene to prevent
back feathers in front and on the sides of the transmitter from covering the solar panel (Fig. 3), that has
noticeably improved data transmission.
Locations of all GPS males were monitored every 3 days during the spring breeding season (~15
Mar – 20 May) in 2014 to determine: (a) locations of potential new leks for crews to check, (b) which
males were alive, (c) which males remained within the study area, (d) which males were attending leks,
(e) inter-lek movements, and (f) nocturnal and diurnal habitat use around leks.
Lek Searching and Lek Counts
Field crews located, verified, and counted 14 previously unknown leks in spring 2014. All but
one of these 14 leks were either found opportunistically during trapping or field work early in the
breeding season or during dual-frame sampling. However, the Tweener lek was discovered by tracking a
yearling to the lek location then later confirmed by a field crew on the ground.
Field crews conducted standard lek counts, unreconciled double-observer counts, and paired
ground and helicopter counts at leks within the study area from 16 March - 22 May 2013. This effort
resulted in data for 93 standard lek counts at 29 different leks, 28 unreconciled double-observer counts at
14 leks, and 46 paired ground and helicopter counts at 21 leks. Field crews entered and proofed all field
data by July 2014.
Survival, Lek Attendance, and Inter-lek Movement
Small sample sizes of birds captured off lek precluded estimating population-level breedingseason survival or lek attendance for GPS males in spring 2013 or spring 2014. Data on inter-lek
movements will be analyzed following the fourth and final field season in spring 2015.
Male Habitat Use Around Leks
We collected breeding-season location data on a total of 31 GPS males in spring 2014. Analysis
of male habitat use around leks from 2012-2014 are underway.
LITERATURE CITED
Aldridge, C. L., S. E. Nielsen, H. L. Beyer, M. S. Boyce, J. W. Connelly, S. T. Knick, and M. A.
Schroeder. 2008. Range-wide patterns of greater sage-grouse persistence. Diversity and Distributions
14:983-994.

13

�Aldridge, C. L., D. J. Saher, T. M. Childers, K. E. Stahlnecker, and Z. H. Bowen. 2011. Crucial nesting
habitat for Gunnison Sage-Grouse: a spatially explicit hierarchical approach. Journal of Wildlife
Management. In press.
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.
Applegate, R. D. 2000. Use and misuse of prairie chicken lek surveys. Wildlife Society Bulletin 28:457458.
Autenrieth, R., W. Molini, and C. E. Braun. 1982. Sage grouse management practices. Western States
Sage Grouse Committee Technical Bulletin 1. Twin Falls, Idaho, USA.
Bartmann, R.M., G.C. White, L.H. Carpenter, and R.A. Garrott. 1987. Aerial mark-recapture estimates of
confined mule deer in pinyon-juniper woodland. Journal of Wildlife Management 51:41-46.
Beck, T. D. I., and C. E. Braun. 1978. Weights of Colorado sage-grouse. Condor 80:241-243.
Beck, T. D. I., and C. E. Braun. 1980. The strutting ground count: variation, traditionalism, and
management needs. Proceedings of the Annual Conference of the Western Association of Fish and
Wildlife Agencies 60:558-566.
Bedrosian, B., and D. Craighead. 2007. Evaluation of techniques for attaching transmitters to common
raven nestlings. Northwestern Naturalist 88:1-6.
Boyko, A. R., R. M. Gibson, and J. R. Lucas. 2004. How predation risk affects the temporal dynamics of
avian leks: greater sage-grouse versus golden eagles. American Naturalist 163: 154-165.
Bowden, D. C., and R. C. Kufeld. 1995. Generalized mark-sight population size estimation applied to
Colorado moose. Journal of Wildlife Management 59:840-851.
Bradbury, J. W., S. L. Verhencamp, and R. M. Gibson. 1989. Dispersion of displaying male sage grouse:
patterns of temporal variation. Behavioral Ecology and Sociobiology 24:1-14.
Braun, C. E., O. O. Oedekoven, and C. L. Aldridge. 2002. Oil and gas development in western North
America: effects on sagebrush steppe avifauna with particular emphasis on sage grouse. Transactions
of the North American Wildlife and Natural Resources Conference 67:337-349.
Colorado Greater Sage-grouse Steering Committee (CGSSC). 2008. Colorado greater sage-grouse
conservation plan. Colorado Division of Wildlife, Denver, CO, USA.
Connelly, J. W., S. T. Knick, M. A. Schroeder, and S. J. Stiver. 2004. Conservation assessment of greater
sage-grouse and sagebrush habitats. Western Association of Fish and Wildlife Agencies, Cheyenne,
Wyoming, USA.
Connelly, J. W., K. P. Reese, and M. A. Schroeder. 2003. Monitoring of greater sage-grouse habitats and
populations. Station Bulletin 80, University of Idaho, Moscow, Idaho, USA.
Connelly, J. W., M. A. Schroeder, A. R. Sands, and C. E. Braun. 2000. Guidelines to manage sage-grouse
populations and their habitats. Wildlife Society Bulletin 28:967-985.
Craighead, D., and B. Bedrosian. 2009. A relationship between blood lead levels of Common Ravens and
the hunting season in the southern Yellowstone Ecosystem. In R. T. Watson, M. Fuller, M. Pokras,
and W. G. Hunt (Eds.). Ingestion of Lead from Spent Ammunition: Implications for Wildlife and
Humans. The Peregrine Fund, Boise, Idaho, USA.
Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. F. Schlatterer. 1963. Ecology,
productivity, and management of sage grouse in Idaho. Journal of Wildlife Management 27:811-840.
Doherty, K. E., D. E. Naugle, H. Copeland, A. Pocewicz, and J. Kiesecker. 2010a. Energy development
and conservation tradeoffs: systematic planning for greater sage-grouse in their eastern range. Studies
in Avian Biology. In press.
Doherty, K. E., D. E. Naugle, and J. Evans. 2010b. A currency for offsetting energy development
impacts: horse-trading sage-grouse on the open market. PLoS ONE 5(4): e10339.
doi:10.1371/journal.pone.0010339.
Doherty, K. E., D. E. Naugle, and B. L. Walker. 2010c. Greater sage-grouse nesting habitat: the
importance of managing at multiple scales. Journal of Wildlife Management 74:1544-1553.

14

�Dunn, P. O., and C. E. Braun. 1985. Natal dispersion and lek fidelity of greater sage-grouse. Auk
102:621-627.
Ellis, K. L. 1984. Behavior of lekking sage-grouse in response to a perched Golden Eagle. Western Birds
15:37-38.
Emmons, S. R., and C. E. Braun. 1984. Lek attendance of male sage grouse. Journal of Wildlife
Management 48:1023–1028.
Fedy, B. C. and C. L. Aldridge. 2011. The importance of within-year repeated counts and the influence of
scale on long-term monitoring of sage-grouse. Journal of Wildlife Management 75: 1022-1033.
Fedy, B. C., C. L. Aldridge, K. E. Doherty, J. L. Beck, B. Bedrosian, M. J. Holloran, G. D. Johnson, N.
W. Kaczor, C. P. Kirol, C. A. Mandich, D. Marshall, G. McKee, C. Olson, C. C. Swanson, and B. L.
Walker. 2012. Inter-seasonal movements of greater sage-grouse, migratory behavior, and an
assessment of the core regions concept in Wyoming. Journal of Wildlife Management 76:1062-1071.
Giesen, K. M., T. J. Schoenberg, and C. E. Braun. 1982. Methods for trapping sage grouse in Colorado.
Wildlife Society Bulletin 10:224-231.
Gill, R. B. 1965. Distribution and abundance of a population of sage grouse in North Park, Colorado.
Thesis, Colorado State University, Fort Collins, Colorado, USA.
Gregg, M. A., J. K. Barnett, and J. A. Crawford. 2008. Temporal variation in diet and nutrition of
preincubating greater sage-grouse. Rangeland Ecology and Management 61:535–542.
Hagen, C. 2010. Greater sage-grouse conservation assessment and strategy for Oregon: a plan to maintain
and enhance populations and habitat. Unpublished report. Oregon Department of Fish and Wildlife.
184 p.
Haines, D. E. and K. H. Pollock. 1998. Estimating the number of active and successful bald eagle nests:
an application of the dual frame method. Environmental and Ecological Statistics 5: 245-256.
Harju, S. M., M. R. Dzialak, R. C. Taylor, L. D. Hayden-Wing, and J. B. Winstead. 2010. Thresholds and
time lags in the effects of energy development on greater sage-grouse populations. Journal of Wildlife
Management 74:437-448.
Hartzler, J. E. 1972. An analysis of sage grouse lek behavior. Ph. D. dissertation. University of Montana,
Missoula, Montana, USA.
Holloran, M. J. 2005. Greater sage-grouse (Centrocercus urophasianus) population response to natural
gas field development in western Wyoming. Ph.D. Dissertation, University of Wyoming, Laramie,
Wyoming, USA.
Holloran, M. J., R. C. Kaiser, and W. A. Hubert. 2010. Yearling greater sage-grouse response to energy
development in Wyoming. Journal of Wildlife Management 74:65-72.
Jenni, D. A., and J. E. Hartzler. 1978. Attendance at a sage grouse lek: implications for spring censuses.
Journal of Wildlife Management 42:46–52.
Krager, R. 1977. Survey of sage grouse strutting ground complexes and seasonal use areas within the
Piceance Basin Wildlife Habitat Area. Unpublished progress report. Colorado Division of Wildlife,
Fort Collins, USA.
Lyon, A. G., and S. H. Anderson. 2003. Potential gas development impacts on sage grouse nest initiation
and movement. Wildlife Society Bulletin 31:486-491.
McClintock, B. T., G. C. White, K. P. Burnham, and M. A. Pryde. 2008. A generalized mixed effects
model of abundance for mark–sight data when sampling is without replacement. Pages 273–292 in D.
L.Thompson, E. G. Cooch, and M. J. Conroy, eds. Modeling demographic processes in marked
populations. Springer, New York, New York, USA.
McRoberts, J. T., M. J. Butler, W. B. Ballard, M. C. Wallace, H. A. Whitlaw, and D. A. Haukos. 2011.
Response of lesser prairie-chickens on leks to aerial surveys. Wildlife Society Bulletin 35:27-31.
Natural Resources Conservation Service (NRCS). 2009. Greater Sage-Grouse Habitat Conservation
Strategy. U. S. Department of Agriculture. Unpublished report. Billings, Montana, USA. 34 p.
Naugle, D. E., and B. L. Walker. 2007. A collaborative vision for integrated monitoring of greater sagegrouse populations. Pages 57-62 in K. P. Reese and T. R. Bowyer, eds. Monitoring populations of

15

�greater sage-grouse: proceedings of a symposium at Idaho State University. College of Natural
Resources Experiment Station Bulletin 88. Moscow, ID.
Neubaum, D. 2011. Parachute-Piceance-Roan Greater Sage-Grouse Newsletter, Vol. 1(1). Unpublished
report, Colorado Division of Wildlife, Grand Junction, USA. 4 pages.
Nichols, J. T., J. E. Hines, J. R. Sauer, F. W. Fallon, J. E. Fallon, and P. J. Heglund. 2000. A doubleobserver approach for estimating detection probability and abundance from point counts. Auk 117:
393-408.
Patterson, R. L. 1952. The sage grouse in Wyoming. Sage Books, Denver, CO USA. 339 p.
Parachute-Piceance-Roan Greater Sage-Grouse Work Group (PPR-GSGWG). 2008. Parachute-PiceanceRoan (PPR) Greater Sage-Grouse Conservation Plan. Colorado Division of Wildlife, Denver,
Colorado.
R Development Core Team 2010. R: a language and environment for statistical computing. Version
2.11.0. R Foundation for Statistical Computing, Vienna, Austria. http://www.R-project.org.
Riddle, J. D., K. H. Pollock, and T. R. Simons. 2010. An unreconciled double-observer method for
estimating detection probability and abundance. Auk 127: 841-849.
Rogers, G. E. 1964. Sage grouse investigations in Colorado. Colorado Game, Fish and Parks Department
Technical Publication Number 16.
Royle, J. A. 2004. N-mixture models for estimating population size from spatially replicated counts.
Biometrics 60:108-115.
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., and C. E. Braun. 1991. Walk-in traps for capturing greater prairie-chickens on leks.
Journal of Field Ornithology 62:378-385.
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.
State of Wyoming. 2010. State of Wyoming Executive Department, Executive Order Number 2010-4.
Office of the Governor. Cheyenne, Wyoming. 15 p.
Tack, J. D. 2010. Sage-grouse and the human footprint: implications for conservation of small and
declining populations. Master’s thesis. University of Montana, Missoula, MT. 96 p.
United States Fish and Wildlife Service (USFWS). 2010. 12-month finding for petitions to list the greater
sage-grouse (Centrocercus urophasianus) as threatened or endangered. Federal Register
75(55):13909-14014.
Wakkinen, W. L., K. P. Reese, J. W. Connelly, and R. A. Fischer. 1992. An improved spotlighting
technique for capturing sage-grouse. Wildlife Society Bulletin 20:425-426.
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. 28 p.
Walker, B. L., D. E. Naugle, and K. E. Doherty. 2007. Greater sage-grouse population response to energy
development and habitat loss. Journal of Wildlife Management 71:2644-2654.
Wallestad, R. and P. Schladweiler. 1974. Breeding season movements and habitat selection of male sage
grouse. Journal of Wildlife Management 38:634-637.
Walsh, D. P., G. C. White, T. E. Remington, and D. C. Bowden. 2004. Evaluation of the lek-count index
for greater sage-grouse. Wildlife Society Bulletin 32:56-68.
Walsh, D. P., J. R. Stiver, G. C. White, T. E. Remington, and A. D. Apa. 2010. Population estimation
techniques for lekking species. Journal of Wildlife Management 74:1607-1613.
Western Association of Fish and Wildlife Agencies (WAFWA). 2008. Greater sage-grouse population
trends: an analysis of lek count databases 1965-2007. Sage- and Columbian Sharp-tailed Grouse
Technical Committee. Unpublished report. Cheyenne, Wyoming, USA. 138 p.
White, G. C. 2010. MARK, Version 6.0. http://warnercnr.colostate.edu/~gwhite/mark/mark.htm.

16

�Zablan, M. A., C. E. Braun, and G. C. White. 2003. Estimation of greater sage-grouse survival in North
Park, Colorado. Journal of Wildlife Management 67:144-154.

17

�FIGURES

Fig. 1. New or potential greater sage-grouse leks (as of May 2014) discovered during field work from
2012-2014 and the estimated occupied range boundary (as of Feb 2012) in the Parachute-Piceance-Roan
population of northwestern Colorado, USA (including the isolated “Magnolia” section).

18

�Figure 2. Original harness and transmitter design for rump-mounted leg-loop attachment of
solar-powered GPS satellite PTT transmitters to male greater sage-grouse.

19

�Figure 3. Improved harness and transmitter design for rump-mounted leg-loop attachment of
solar-powered GPS satellite PTT transmitters for male greater sage-grouse. This photo also
shows the underside placement of a micro-VHF mortality sensor/transmitter.

20

�Figure 4. Attachment, placement, and camouflage of rump-mounted, solar-powered, GPS
satellite PTT transmitters for male greater sage-grouse.

21

�Figure 5. Drop net design for capturing male greater sage-grouse.

22

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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
Evaluating Lek-Based Monitoring and
Management Strategies for Greater Sage-Grouse in
the Parachute-Piceance-Roan Population in
Northwestern Colorado

Period Covered: September 1, 2014 – August 31, 2015
Author: B. L. Walker
Personnel: B. Holmes, S. Duckett, B. Petch, W. deVergie, J. T. Romatzke
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
Implementing effective monitoring and mitigation strategies is crucial for conserving populations of
sensitive wildlife species. Concern over the status of greater sage-grouse populations has increased rangewide and in Colorado due to population declines, range contraction, loss and degradation of sagebrush
habitat, and potential for listing the species under the Endangered Species Act. Despite untested
assumptions, lek counts are widely used as an index of abundance by state agencies to monitor sagegrouse populations. Lek locations are also commonly used to identify and protect important sage-grouse
habitat. However, the use of lek counts and locations to monitor and manage sage-grouse populations
remains controversial because it is unknown how closely lek-count data track actual changes in male
abundance from year to year or if lek buffers are effective at protecting habitat for male sage-grouse
during the breeding season. Colorado Parks and Wildlife deployed solar-powered GPS satellite
transmitters on male greater sage-grouse to obtain data on male survival, lek attendance, inter-lek
movements, and diurnal and nocturnal habitat use around leks and conducted double-observer lek counts
to estimate detectability of males on leks during the breeding season in the Parachute-Piceance-Roan
population in northwestern Colorado in spring from 2012-2015. These data will allow us to evaluate
whether current lek-based monitoring methods provide reliable information about sage-grouse population
trends and whether current lek buffers are effective at protecting breeding males. Chevron did not provide
access in spring 2015, so no additional GPS males were marked. We monitored 17 GPS males for part or
all of the 1 September 2014 - 31 August 2015 period. Analyses for this project are in progress.

1

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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
Evaluating Lek-Based Monitoring and
Management Strategies for Greater Sage-Grouse in
the Parachute-Piceance-Roan Population in
Northwestern Colorado

Period Covered: September 1, 2015 – August 31, 2016
Author: B. L. Walker
Personnel: B. Holmes, S. Duckett, B. Petch, B. deVergie, J. T. Romatzke
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
Implementing effective monitoring and mitigation strategies is crucial for conserving populations of
sensitive wildlife species. Concern over the status of greater sage-grouse populations has increased rangewide and in Colorado due to population declines, range contraction, loss and degradation of sagebrush
habitat, and potential for listing the species under the Endangered Species Act. Despite untested
assumptions, lek counts are widely used as an index of abundance by state agencies to monitor sagegrouse populations. Lek locations are also commonly used to identify and protect important sage-grouse
habitat. However, the use of lek counts and locations to monitor and manage sage-grouse populations
remains controversial because it is unknown how closely lek-count data track actual changes in male
abundance from year to year or if lek buffers are effective at protecting habitat for male sage-grouse
during the breeding season. Colorado Parks and Wildlife deployed solar-powered GPS satellite
transmitters on male greater sage-grouse to obtain data on male survival, lek attendance, inter-lek
movements, and diurnal and nocturnal habitat use around leks and conducted double-observer lek counts
to estimate detectability of males on leks during the breeding season in the Parachute-Piceance-Roan
population in northwestern Colorado in spring from 2012-2016. These data will allow us to evaluate
whether current lek-based monitoring methods provide reliable information about sage-grouse population
trends and whether current lek buffers are effective at protecting breeding males. We monitored 17 GPS
males for all or part of the 1 September 2014 - 31 August 2015 period and 6 males for all or part of the 1
September 2015 – 31 May 2016 period. Analyses for this project are in progress.

1

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