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                  <text>COLORADO PARKS AND WILDLIFE - AVIAN RESEARCH PROGRAM
Progress Report
November 2015
TITLE: Greater Sage-grouse natal dispersal and brood augmentation with captive-reared chicks
AUTHOR: Anthony D. Apa
PROJECT PERSONNEL: T. R. Thompson and K. P. Reese, University of Idaho, Moscow, ID
Period Covered: March 1, 2005 – November 1, 2015
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NOT BE PUBLISHED OR QUOTED without permission of the author.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.
PROJECT ABSTRACT
In response to population declines, recent research on greater sage-grouse (Centrocercus urophasianus)
has focused on the population ecology, habitat relationships, and response to management practices by
this species. However, the mechanisms, patterns, and consequences of movements between seasonal
habitats, especially by juveniles during natal dispersal, and the effects of this movement on survival,
recruitment, the redistribution of individuals, as well as the population dynamics within and between
populations remains largely unknown. Quantifiable data and information on juvenile dispersal and
survival in the greater sage-grouse is one of the least understood aspects of this species’ life history.
Dispersal patterns and recruitment processes of juvenile sage-grouse, as well the landscape characteristics
that influence and contribute to these movements remain lacking. Knowledge of the dispersal ecology
(timing, distances moved, frequency and rate of movement, immigration and emigration rates within and
between populations, and juvenile survivorship) will provide better information on how to manage this
species at the landscape level, as well as within and between populations. This information will be useful
in attempting to improve and plan for the conservation and management of this species as its habitat
becomes more fragmented and altered. The objectives of our study were to 1) determine the sex-specific
movement patterns of juvenile sage-grouse during natal dispersal including timing, duration, rate of
movement, distances moved and recruitment rate, 2) determine the effects of these dispersal patterns on
survival rates and causes of mortality, 3) determine how landscape structure influences both the
movement patterns and survival of juveniles during this period, 4) verify and evaluate the mechanisms
and conditions of adoption in wild broods through the introduction of domestically-hatched chicks and
observation of natural adoption rates, 5) assess the movement patterns and survivorship of successfully
adopted domestically-hatched 2 and 7 day-old chicks from the natal area of the surrogate brood to chick
independence and brood break-up (approximately 10 weeks of age), and 6) compare the movement
patterns and survivorship of domestically-hatched chicks with the movement patterns and survivorship of
wild-hatched chicks in mixed and unmixed broods from the natal area of the surrogate brood to chick
independence and brood break-up. The study areas were located in the Axial Basin and Cold Springs
Mountain in northwestern Colorado from 2005 – 2007. The project field research and final report is
complete and we continue in the publication phase of this research project.

�Objectives 4, 5, and 6
Captive rearing sage-grouse for augmentation of surrogate wild broods: evidence for success
Abstract-Both species of North American sage-grouse (Centrocercus spp.) have experienced declines in
distribution and abundance. Translocation of adult birds from a stable population to a small or declining
population has been a management tool used by wildlife managers to support population persistence in
these areas. Captive rearing chicks and releasing them into wild surrogate broods is an untested
alternative to augment declining populations of sage-grouse. We developed techniques to successfully
rear sage-grouse chicks in captivity, evaluated explanatory variables that could influence hatch and
captive-rearing success, and estimated the survival of domestically hatched (DH) chicks to 28 days of age
following introduction to a surrogate wild brood.We collected 304 eggs from radiomarked female greater
sage-grouse (C. urophasianus) during 2004–2007 in 3 study areas in northwestern Colorado. Estimated
hatching success of collected eggs was 0.745 (SE¼0.022, 95% CI¼0.700–0.786) and was negatively
influenced by the number of days an egg was stored and the percent egg weight loss that occurred during
storage and incubation. We monitored 175 DH chicks in captivity for 1–10 days before introduction and
adoption into surrogate wild broods. Model-averaged captive-rearing success was 0.792 (SE¼0.045, 95%
CI¼0.686–0.865) across years, and was positively influenced by initial chick mass at hatch and daily
weight gain in captivity but negatively influenced by the number of days the egg was stored and
advancing hatch date. We were able to radiomark and monitor 133 DH chicks adopted into surrogate wild
broods until 28 days of age. Eighty-eight percent of DH chicks were successfully adopted within 24
hours. Our overall estimate of DH chick survival to 28 days (0.423; 95% CI¼0.257–0.587) was
comparable to published wild-hatched chick survival. Predation and exposure-related deaths accounted
for 26.3% and 25.6% of the known fates, respectively. Our captive-rearing protocols and techniques were
successful for collecting greater sage-grouse eggs, hatching and rearing chicks in captivity, and releasing
chicks into wild surrogate broods. This success further implies that captive rearing and release can be a
potential management strategy to demographically and genetically reinforce or augment small
populations of sage-grouse. © 2015 The Wildlife Society.
Publication:
Thompson, T. R., A. D. Apa, K. P. Reese, and K. M. Tadvick. 2015. Captive rearing sage-grouse for
augmentation of surrogate wild broods; evidence for success. Journal of Wildlife Management
79:998-1013.
Objectives 1 &amp; 2:
Survival of greater sage-grouse broods and chicks from hatch to brood independence in
northwestern Colorado
Abstract-Survival of chicks from hatch to brood independence and recruitment into fall populations is an
important but poorly understood life history trait that can have important consequences on the dynamics
and viability of greater sage-grouse (Centrocercus urophasianus) populations. Little is known about how
the factors of gender, hatch date, hatch weight, distance traveled from nest, and brood size contribute both
individually and ecologically to survival of chicks. We monitored survival and causes of mortality in
wild-hatched (WH) chicks (n = 431) in wild broods (n = 115) from hatch to 16 weeks of age in the AB
(Axial Basin) and CSM (Cold Springs Mountain) study areas in northwestern Colorado, 2005-2007 and
evaluated potentially important predictors of brood and chick survival. In addition, we monitored
survival from hatch to 16 weeks of age for a cohort of domestically-hatched (DH) chicks raised to 1-10
days of age in captivity (n = 116) and introduced into a subset of wild broods during this same time
period. Overall brood survival from 2005-2007 (both wild and wild broods augmented with DH chicks)

�to 16 weeks of age was 0.381 (95% CI: 0.264 – 0.514) at CSM compared to 0.533 (0.405 – 0.657) in the
AB. Within the AB, we observed higher model-averaged survival rates among broods with DH chicks
(0.631, SE = 0.088) compared to broods without (0.430, SE = 0.104), while at CSM the pattern was
reversed (0.205, SE = 0.102 and 0.573, SE = 0.080). When we included broods that were depredated 1-3
days post-hatch and before radiomarking of chicks our overall apparent brood survival decreased from
47.8% (55/115) to 43.7% (55/126). The main cause of chick death was from predation, although
exposure accounted for 27% of mortalities among DH chicks. Model averaged estimates of brood and
chick survival indicated that survival varied both temporally and spatially. Brood and chick survival were
higher in the AB compared to CSM, and WH chicks had higher survival in both areas compared to DH
chicks. Similarly, DH and WH chicks at CSM in both augmented and wild broods had lower survival
than DH and WH chicks in the AB with the largest differences occurring during weeks 1 – 3. Among the
WH chicks survival rates among years in the AB ranged from 0.158 to 0.446 and at CSM from 0.088 to
0.339. Between study areas and among years survival was lowest during the first 3 – 4 weeks. We found
evidence that chick survival increased with age and decreased with advancing hatch date, but found
limited support for the influence of gender or distance traveled. We recommend that managers develop
better understanding and knowledge of the relationship between nesting cover and brood habitat, as well
as movement patterns between these areas within a landscape for each population. Managers need to
consider prioritizing the protection and restoration of both early- and late brood-rearing habitat within
specific landscapes, as our study demonstrates 2 bottlenecks through which chick survival significantly
decrease at &lt; 21-day post-hatch and during brood independence at &gt; 10 weeks of age. We suggest that &gt;
3 areas of each seasonal brood habitat type be dispersed within a breeding population to maintain
traditional use patterns and to facilitate the use of new areas (i.e., restorations or plantings such as CRP),
so as to help reduce predation risks and exposures due to concentration of broods in poor quality or
limited critical habitat.
Publication:
Thompson, T. R., A. D. Apa, and K. P. Reese. In Prep. Survival of greater sage-grouse broods and chicks
from hatch to brood independence in northwestern Colorado. Journal of Wildlife Management
XXX:XXXX-XXXX.
Objectives 1 &amp; 2:
Survival, natal dispersal and recruitment of juvenile greater sage-grouse in northwest Colorado
Abstract-Juvenile survival and recruitment has not been studied extensively in many grouse species,
including greater sage-grouse (Centrocercus urophasianus). Since there is scant information on this vital
rate, the implications of management actions on specific population demographics remains unknown. We
captured, radio-marked, and monitored survival and recruitment of 183 transmitter-equipped juvenile
sage-grouse from 1 September – 31 March at 2 study areas in northwest Colorado (AB: Axial Basin,
CMS: Cold Springs Mountain). Juvenile grouse survival September through March varied by month,
study area, and gender. Juvenile females had higher survival than juvenile males, and survival for each
was higher in AB compared to CSM. Juvenile survival was lowest during September and October and
coincided with brood independence and integration into winter flocks. Average survival from hatch to
recruitment into the natal breeding population (March) varied between areas (AB: x̄ = 0.287, SE = 0.039;
CSM: x̄ = 0.122, SE = 0.054). This information on survival and recruitment of juvenile sage-grouse has
important implications for the management of this species at local, landscape, and regional levels.

�Publication:
Apa, A. D., T. R. Thompson, and K. P. Reese. In Prep. Juvenile greater sage-grouse survival, fall
movements, and recruitment in northwest Colorado. Journal of Wildlife Management
XXX:XXXX-XXXX.
Objectives 1 &amp; 2:
Dispersal, gene flow, and population genetic structure in the greater sage-grouse: implications for
connectivity and natural recolonization

Abstract. Dispersal and gene flow have important consequences for the population dynamics
and genetic structure of populations. However, for most species the degree to which dispersal
and gene flow maintain population demographic and genetic connectivity remains unknown.
Here, we compare the patterns of dispersal and genetic structure in Greater Sage-Grouse
(Centrocercus urophasianus) at 15 leks in six population management zones (PMZs) in
northwest Colorado by genotyping 275 individuals at 17 microsatellite loci. All leks showed
high levels of genetic diversity, and low levels of genetic structure were observed between
neighboring leks and PMZs. Multiple analyses revealed an isolation by distance pattern among
leks and PMZs that followed a directional or two-dimensional stepping-stone pattern. Contrary
to the traditional view of female-biased dispersal in avian and grouse species, we observed direct
evidence of male-biased dispersal in our radio-telemetry data, but less evidence in our genetic
methods. Our spatial autocorrelation analyses revealed significantly positive r values out to the
5 km distance class for males and 15 km distance class for females. Analyses of dispersal using
direct and indirect methods indicated that dispersal distances above 20 km are rare. Our study
demonstrates the importance of using both demographic and genetic methods to define and
understand population characteristics and manage populations at appropriate scales.
Publication:
Apa, A. D. T. R. Thompson, S. Oyler-McCance, K. P. Reese, and L. P. Waits. In Revision. Comparing
dispersal and gene flow in the greater sage-grouse: implications for connectivity and natural
recolonizaton. Condor XXX:XXXX-XXXX.

Objective 3:
Relationship of landscape characteristics to movement behaviors and settlement patterns of greater
sage-grouse in northwest Colorado
Abstract-Range-wide declines in greater sage-grouse (Centrocercus urophasiansus) populations have
largely been attributed to loss, degradation, and fragmentation of sagebrush habitats and landscapes that
are believed to negatively impact population vital rates, movements, and distribution patterns. Current
understanding of these processes in sage-grouse is primarily limited to adult age individuals with little
understanding of their influences on juvenile movement behaviors and settlement patterns. In this study
we assessed how landscape composition (percent land cover) and edge density (m/ ha) within the
dispersal range (winter and dispersal locations) and dispersal period landscapes (pre-dispersal, winter, and
post-dispersal locations) differed between male and female juvenile sage-grouse in 2 study areas (Axial
Basin and Cold Springs Mountain) in northwestern Colorado. During September – April, 2005 – 2008 we

�monitored 95 juveniles (74 female and 31 males). Before running landscape analyses we performed an
accuracy assessment on 3 potential Landsat satellite imagery sources (Colorado Vegetation Classification
Project, LANDFIRE, and Southwest Regional GAP) and used overall accuracy, and kappa coefficients to
determine which data source would have the highest quality and less uncertainty in derived land cover
maps. Using the LANDFIRE (2006) Existing Vegetation Map we compared proportion of land cover
types and edge densities in 4 dominant land cover types (sagebrush dominated community (Artemisia
tridentata spp), salt desert shrub dominated community (shadscale saltbush (Atriplex confertifolia);
greasewood (Sarcobatus vermiculatus)), grassland/ rangeland/ perennial grass and forb, and deciduous
shrub/ mountain-shrub dominated community (bitterbrush (Purshia tridentate); Gambel oak (Quercus
gambelii); serviceberry (Amelanchier spp); snowberry (Symphoricarpos spp.), and tested for effect on
genders, areas, dispersal ranges, and among dispersal period landscapes at 2 spatial extents (500- and
2,000-m). Dispersal ranges and dispersal period landscape metrics were not significantly different
between genders at either buffer extent. Within dispersal ranges, percent cover in sagebrush did not
significantly differ between study areas at the 500-m buffer extent; however at the 2,000-m buffer extent
proportion of land cover in sagebrush was higher in the Axial Basin. Among dispersal period landscapes,
measured metrics significantly differed between areas and among periods. At the 500-m buffer extent
winter and post-dispersal landscapes in the Axial Basin had higher land cover in sagebrush, lower edge
density in sagebrush, and lower cover in salt desert shrub compared to Cold Springs Mountain. At the
2,000-m buffer extent a similar pattern was observed, as well as higher land cover in sagebrush and shrub,
as well as shrub edge density in the Axial Basin. The grassland cover type did not significantly differ at
either buffer extent for dispersal range or dispersal period landscapes. We believe this suggests natal
dispersal movement behaviors and settlement patterns within our study areas, where percent land cover in
sagebrush are &gt; 60%, are not directly influenced by landscape structure or composition in the dispersal
range or period, but by individual and population pressures and demands (e.g., access to resources,
inbreeding avoidance, traditional use) related to the breeding and production (brood-rearing) areas.

Publication:
Thompson, T. R., A. D. Apa, and K. P. Reese. In Prep. Relationship of landscape characteristics to
movement behaviors and settlement patterns of greater sage-grouse in northwest Colorado.
Journal of Wildlife Management XXX:XXXX-XXXX.

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              <text>In response to population declines, recent research on greater sage-grouse (&lt;em&gt;Centrocercus urophasianus&lt;/em&gt;) has focused on the population ecology, habitat relationships, and response to management practices by this species. However, the mechanisms, patterns, and consequences of movements between seasonal habitats, especially by juveniles during natal dispersal, and the effects of this movement on survival, recruitment, the redistribution of individuals, as well as the population dynamics within and between populations remains largely unknown. Quantifiable data and information on juvenile dispersal and survival in the greater sage-grouse is one of the least understood aspects of this species’ life history. Dispersal patterns and recruitment processes of juvenile sage-grouse, as well the landscape characteristics that influence and contribute to these movements remain lacking. Knowledge of the dispersal ecology (timing, distances moved, frequency and rate of movement, immigration and emigration rates within and between populations, and juvenile survivorship) will provide better information on how to manage this species at the landscape level, as well as within and between populations. This information will be useful in attempting to improve and plan for the conservation and management of this species as its habitat becomes more fragmented and altered. The objectives of our study were to 1) determine the sex-specific movement patterns of juvenile sage-grouse during natal dispersal including timing, duration, rate of movement, distances moved and recruitment rate, 2) determine the effects of these dispersal patterns on survival rates and causes of mortality, 3) determine how landscape structure influences both the movement patterns and survival of juveniles during this period, 4) verify and evaluate the mechanisms and conditions of adoption in wild broods through the introduction of domestically-hatched chicks and observation of natural adoption rates, 5) assess the movement patterns and survivorship of successfully adopted domestically-hatched 2 and 7 day-old chicks from the natal area of the surrogate brood to chick independence and brood break-up (approximately 10 weeks of age), and 6) compare the movement patterns and survivorship of domestically-hatched chicks with the movement patterns and survivorship of wild-hatched chicks in mixed and unmixed broods from the natal area of the surrogate brood to chick independence and brood break-up. The study areas were located in the Axial Basin and Cold Springs Mountain in northwestern Colorado from 2005 – 2007. The project field research and final report is complete and we continue in the publication phase of this research project.</text>
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