Phenology largely explains taller grass at successful nests in greater sage‐grouse

Much interest lies in the identification of manageable habitat variables that affect key vital rates for species of concern. For ground‐nesting birds, vegetation surrounding the nest may play an important role in mediating nest success by providing concealment from predators. Height of grasses surrounding the nest is thought to be a driver of nest survival in greater sage‐grouse (Centrocercus urophasianus; sage‐grouse), a species that has experienced widespread population declines throughout their range. However, a growing body of the literature has found that widely used field methods can produce misleading inference on the relationship between grass height and nest success. Specifically, it has been demonstrated that measuring concealment following nest fate (failure or hatch) introduces a temporal bias whereby successful nests are measured later in the season, on average, than failed nests. This sampling bias can produce inference suggesting a positive effect of grass height on nest survival, though the relationship arises due to the confounding effect of plant phenology, not an effect on predation risk. To test the generality of this finding for sage‐grouse, we reanalyzed existing datasets comprising >800 sage‐grouse nests from three independent studies across the range where there was a positive relationship found between grass height and nest survival, including two using methods now known to be biased. Correcting for phenology produced equivocal relationships between grass height and sage‐grouse nest survival. Viewed in total, evidence for a ubiquitous biological effect of grass height on sage‐grouse nest success across time and space is lacking. In light of these findings, a reevaluation of land management guidelines emphasizing specific grass height targets to promote nest success may be merited.     

Predicting greater sage‐grouse habitat selection at the southern periphery of their range

Mapping suitable habitat is an important process in wildlife conservation planning. Species distribution reflects habitat selection processes occurring across multiple spatio‐temporal scales. Because habitat selection may be driven by different factors at different scales, conservation planners require information at the scale of the intervention to plan effective management actions. Previous research has described habitat selection processes shaping the distribution of greater sage‐grouse (Centrocercus urophasianus; sage‐grouse) at the range‐wide scale. Finer‐scale information for applications within jurisdictional units inside the species range is lacking, yet necessary, because state wildlife agencies are the management authority for sage‐grouse in the United States. We quantified seasonal second‐order habitat selection for sage‐grouse across the state of Utah to produce spatio‐temporal predictions of their distribution at the southern periphery of the species range. We used location data obtained from sage‐grouse marked with very‐high‐frequency radio‐transmitters and lek location data collected between 1998 and 2013 to quantify species habitat selection in relation to a suite of topographic, edaphic, climatic, and anthropogenic variables using random forest algorithms. Sage‐grouse selected for greater sagebrush (Artemisia spp.) cover, higher elevations, and gentler slopes and avoided lower precipitations and higher temperatures. The strength of responses to habitat variables varied across seasons. Anthropogenic variables previously reported as affecting their range‐wide distribution (i.e., roads, powerlines, communication towers, and agricultural development) were not ranked as top predictors at our focal scale. Other than strong selection for sagebrush cover, the responses we observed differed from what has been reported at the range‐wide scale. These differences likely reflect the unique climatic, geographic, and topographic context found in the southern peripheral area of the species distribution compared to range‐wide environmental gradients. Our results highlight the importance of considering appropriateness of scale when planning conservation actions for wide‐ranging species.     

Modeling ecological minimum requirements for distribution of greater sage‐grouse leks: implications for population connectivity across their western range,U.S.A

Greater sage‐grouse Centrocercus urophasianus (Bonaparte) currently occupy approximately half of their historical distribution across western North America. Sage‐grouse are a candidate for endangered species listing due to habitat and population fragmentation coupled with inadequate regulation to control development in critical areas. Conservation planning would benefit from accurate maps delineating required habitats and movement corridors. However, developing a species distribution model that incorporates the diversity of habitats used by sage‐grouse across their widespread distribution has statistical and logistical challenges. We first identified the ecological minimums limiting sage‐grouse, mapped similarity to the multivariate set of minimums, and delineated connectivity across a 920,000 km² region. We partitioned a Mahalanobis D² model of habitat use into k separate additive components each representing independent combinations of species–habitat relationships to identify the ecological minimums required by sage‐grouse. We constructed the model from abiotic, land cover, and anthropogenic variables measured at leks (breeding) and surrounding areas within 5 km. We evaluated model partitions using a random subset of leks and historic locations and selected D² (k = 10) for mapping a habitat similarity index (HSI). Finally, we delineated connectivity by converting the mapped HSI to a resistance surface. Sage‐grouse required sagebrush‐dominated landscapes containing minimal levels of human land use. Sage‐grouse used relatively arid regions characterized by shallow slopes, even terrain, and low amounts of forest, grassland, and agriculture in the surrounding landscape. Most populations were interconnected although several outlying populations were isolated because of distance or lack of habitat corridors for exchange. Land management agencies currently are revising land‐use plans and designating critical habitat to conserve sage‐grouse and avoid endangered species listing. Our results identifying attributes important for delineating habitats or modeling connectivity will facilitate conservation and management of landscapes important for supporting current and future sage‐grouse populations.     

The genetic network of greater sage‐grouse: Range‐wide identification of keystone hubs of connectivity

Genetic networks can characterize complex genetic relationships among groups of individuals, which can be used to rank nodes most important to the overall connectivity of the system. Ranking allows scarce resources to be guided toward nodes integral to connectivity. The greater sage‐grouse (Centrocercus urophasianus) is a species of conservation concern that breeds on spatially discrete leks that must remain connected by genetic exchange for population persistence. We genotyped 5,950 individuals from 1,200 greater sage‐grouse leks distributed across the entire species’ geographic range. We found a small‐world network composed of 458 nodes connected by 14,481 edges. This network was composed of hubs—that is, nodes facilitating gene flow across the network—and spokes—that is, nodes where connectivity is served by hubs. It is within these hubs that the greatest genetic diversity was housed. Using indices of network centrality, we identified hub nodes of greatest conservation importance. We also identified keystone nodes with elevated centrality despite low local population size. Hub and keystone nodes were found across the entire species’ contiguous range, although nodes with elevated importance to network‐wide connectivity were found more central: especially in northeastern, central, and southwestern Wyoming and eastern Idaho. Nodes among which genes are most readily exchanged were mostly located in Montana and northern Wyoming, as well as Utah and eastern Nevada. The loss of hub or keystone nodes could lead to the disintegration of the network into smaller, isolated subnetworks. Protecting both hub nodes and keystone nodes will conserve genetic diversity and should maintain network connections to ensure a resilient and viable population over time. Our analysis shows that network models can be used to model gene flow, offering insights into its pattern and process, with application to prioritizing landscapes for conservation.     

Sagebrush treatments influence annual population change for greater sage‐grouse

Vegetation management practices have been applied worldwide to enhance habitats for a variety of wildlife species. Big sagebrush (Artemisia tridentata spp.) communities, iconic to western North America, have been treated to restore herbaceous understories through chemical, mechanical, and prescribed burning practices thought to improve habitat conditions for greater sage‐grouse (Centrocercus urophasianus) and other species. Although the response of structural attributes of sagebrush communities to treatments is well understood, there is a need to identify how treatments influence wildlife population dynamics. We investigated the influence of vegetation treatments occurring in Wyoming, United States, from 1994 to 2012 on annual sage‐grouse population change using yearly male sage‐grouse lek counts. We investigated this response across 1, 3, 5, and 10‐year post‐treatment lags to evaluate how the amount of treated sagebrush communities and time since treatment influenced population change, while accounting for climate, wildfire, and anthropogenic factors. With the exception of chemical treatments exhibiting a positive association with sage‐grouse population change 11 years after implementation, population response to treatments was either neutral or negative for at least 11 years following treatments. Our work supports a growing body of research advocating against treating big sagebrush habitats for sage‐grouse, particularly in Wyoming big sagebrush (A. t. wyomingensis). Loss and fragmentation of sagebrush habitats has been identified as a significant threat for remaining sage‐grouse populations. Because sagebrush may take decades to recover following treatments, we recommend practitioners use caution when designing projects to alter remaining habitats, especially when focused on habitat requirements for one life stage and a single species.     

Landscape characteristics influencing the genetic structure of greater sage‐grouse within the stronghold of their range: a holistic modeling approach

Given the significance of animal dispersal to population dynamics and geographic variability, understanding how dispersal is impacted by landscape patterns has major ecological and conservation importance. Speaking to the importance of dispersal, the use of linear mixed models to compare genetic differentiation with pairwise resistance derived from landscape resistance surfaces has presented new opportunities to disentangle the menagerie of factors behind effective dispersal across a given landscape. Here, we combine these approaches with novel resistance surface parameterization to determine how the distribution of high‐ and low‐quality seasonal habitat and individual landscape components shape patterns of gene flow for the greater sage‐grouse (Centrocercus urophasianus) across Wyoming. We found that pairwise resistance derived from the distribution of low‐quality nesting and winter, but not summer, seasonal habitat had the strongest correlation with genetic differentiation. Although the patterns were not as strong as with habitat distribution, multivariate models with sagebrush cover and landscape ruggedness or forest cover and ruggedness similarly had a much stronger fit with genetic differentiation than an undifferentiated landscape. In most cases, landscape resistance surfaces transformed with 17.33‐km‐diameter moving windows were preferred, suggesting small‐scale differences in habitat were unimportant at this large spatial extent. Despite the emergence of these overall patterns, there were differences in the selection of top models depending on the model selection criteria, suggesting research into the most appropriate criteria for landscape genetics is required. Overall, our results highlight the importance of differences in seasonal habitat preferences to patterns of gene flow and suggest the combination of habitat suitability modeling and linear mixed models with our resistance parameterization is a powerful approach to discerning the effects of landscape on gene flow.     

Greater sage‐grouse respond positively to intensive post‐fire restoration treatments

Habitat loss is the most prevalent threat to biodiversity in North America. One of the most threatened landscapes in the United States is the sagebrush (Artemisia spp.) ecosystem, much of which has been fragmented or converted to non‐native grasslands via the cheatgrass‐fire cycle. Like many sagebrush obligates, greater sage‐grouse (Centrocercus urophasianus) depend upon sagebrush for food and cover and are affected by changes to this ecosystem. We investigated habitat selection by 28 male greater sage‐grouse during each of 3 years after a 113,000‐ha wildfire in a sagebrush steppe ecosystem in Idaho and Oregon. During the study period, seeding and herbicide treatments were applied for habitat restoration. We evaluated sage‐grouse responses to vegetation and post‐fire restoration treatments. Throughout the 3 years post‐fire, sage‐grouse avoided areas with high exotic annual grass cover but selected strongly for recovering sagebrush and moderately strongly for perennial grasses. By the third year post‐fire, they preferred high‐density sagebrush, especially in winter when sagebrush is the primary component of the sage‐grouse diet. Sage‐grouse preferred forb habitat immediately post‐fire, especially in summer, but this selection preference was less strong in later years. They also selected areas that were intensively treated with herbicide and seeded with sagebrush, grasses, and forbs, although these responses varied with time since treatment. Wildfire can have severe consequences for sagebrush‐obligate species due to loss of large sagebrush plants used for food and for protection from predators and thermal extremes. Our results show that management efforts, including herbicide application and seeding of plants, directed at controlling exotic annual grasses after a wildfire can positively affect habitat selection by sage‐grouse.     

A method for estimating population sex ratio for sage‐grouse using noninvasive genetic samples

Population sex ratio is an important metric for wildlife management and conservation, but estimates can be difficult to obtain, particularly for sexually monomorphic species or for species that differ in detection probability between the sexes. Noninvasive genetic sampling (NGS) using polymerase chain reaction (PCR) has become a common method for identifying sex from sources such as hair, feathers or faeces, and is a potential source for estimating sex ratio. If, however, PCR success is sex‐biased, naively using NGS could lead to a biased sex ratio estimator. We measured PCR success rates and error rates for amplifying the W and Z chromosomes from greater sage‐grouse (Centrocercus urophasianus) faecal samples, examined how success and error rates for sex identification changed in response to faecal sample exposure time, and used simulation models to evaluate precision and bias of three sex assignment criteria for estimating population sex ratio with variable sample sizes and levels of PCR replication. We found PCR success rates were higher for females than males and that choice of sex assignment criteria influenced the bias and precision of corresponding sex ratio estimates. Our simulations demonstrate the importance of considering the interplay between the sex bias of PCR success, number of genotyping replicates, sample size, true population sex ratio and accuracy of assignment rules for designing future studies. Our results suggest that using faecal DNA for estimating the sex ratio of sage‐grouse populations has great potential and, with minor adaptations and similar marker evaluations, should be applicable to numerous species.     

Nesting,brood rearing,and summer habitat selection by translocated greater sage‐grouse in North Dakota,USA

Human enterprise has led to large‐scale changes in landscapes and altered wildlife population distribution and abundance, necessitating efficient and effective conservation strategies for impacted species. Greater sage‐grouse (Centrocercus urophasianus; hereafter sage‐grouse) are a widespread sagebrush (Artemisia spp.) obligate species that has experienced population declines since the mid‐1900s resulting from habitat loss and expansion of anthropogenic features into sagebrush ecosystems. Habitat loss is especially evident in North Dakota, USA, on the northeastern fringe of sage‐grouse’ distribution, where a remnant population remains despite recent development of energy‐related infrastructure. Resource managers in this region have determined a need to augment sage‐grouse populations using translocation techniques that can be important management tools for countering species decline from range contraction. Although translocations are a common tool for wildlife management, very little research has evaluated habitat following translocation, to track individual behaviors such as habitat selection and fidelity to the release site, which can help inform habitat requirements to guide selection of future release sites. We provide an example where locations from previously released radio‐marked sage‐grouse are used in a resource selection function framework to evaluate habitat selection following translocation and identify areas of seasonal habitat to inform habitat management and potential restoration needs. We also evaluated possible changes in seasonal habitat since the late 1980s using spatial data provided by the Rangeland Analysis Platform coupled with resource selection modeling results. Our results serve as critical baseline information for habitat used by translocated individuals across life stages in this study area, and will inform future evaluations of population performance and potential for long‐term recovery.     

Spatially explicit models of seasonal habitat for greater sage‐grouse at broad spatial scales: Informing areas for management in Nevada and northeastern California

Defining boundaries of species' habitat across broad spatial scales is often necessary for management decisions, and yet challenging for species that demonstrate differential variation in seasonal habitat use. Spatially explicit indices that incorporate temporal shifts in selection can help overcome such challenges, especially for species of high conservation concern. Greater sage‐grouse Centrocercus urophasianus (hereafter, sage‐grouse), a sagebrush obligate species inhabiting the American West, represents an important case study because sage‐grouse exhibit seasonal habitat patterns, populations are declining in most portions of their range and are central to contemporary national land use policies. Here, we modeled spatiotemporal selection patterns for telemetered sage‐grouse across multiple study sites (1,084 sage‐grouse; 30,690 locations) in the Great Basin. We developed broad‐scale spatially explicit habitat indices that elucidated space use patterns (spring, summer/fall, and winter) and accounted for regional climatic variation using previously published hydrographic boundaries. We then evaluated differences in selection/avoidance of each habitat characteristic between seasons and hydrographic regions. Most notably, sage‐grouse consistently selected areas dominated by sagebrush with few or no conifers but varied in type of sagebrush selected by season and region. Spatiotemporal variation was most apparent based on availability of water resources and herbaceous cover, where sage‐grouse strongly selected upland natural springs in xeric regions but selected larger wet meadows in mesic regions. Additionally, during the breeding period in spring, herbaceous cover was selected strongly in the mesic regions. Lastly, we expanded upon an existing joint–index framework by combining seasonal habitat indices with a probabilistic index of sage‐grouse abundance and space use to produce habitat maps useful for sage‐grouse management. These products can serve as conservation planning tools that help predict expected benefits of restoration activities, while highlighting areas most critical to sustaining sage‐grouse populations. Our joint–index framework can be applied to other species that exhibit seasonal shifts in habitat requirements to help better guide conservation actions.     

Restoring Sage‐grouse nesting habitat through removal of early successional conifer

Conifer woodlands have expanded into sagebrush (Artemisia spp.) ecosystems and degrade habitat for sagebrush obligate species such as the Greater Sage‐grouse (Centrocercus urophasianus). Conifer management is increasing despite a lack of empirical evidence assessing outcomes to grouse and their habitat. Although assessments of vegetation recovery after conifer removal are common, comparisons of successional trends with habitat guidelines or actual data on habitat used by sage‐grouse is lacking. We assessed impacts of conifer encroachment on vegetation characteristics known to be important for sage‐grouse nesting. Using a controlled repeated measures design, we then evaluated vegetation changes for 3 years after conifer removal. We compared these results to data from 356 local sage‐grouse nests, rangewide nesting habitat estimates, and published habitat guidelines. We measured negative effects of conifer cover on many characteristics important for sage‐grouse nesting habitat including percent cover of forbs, grasses, and shrubs, and species richness of forbs and shrubs. In untreated habitat, herbaceous vegetation cover was slightly below the cover at local nest sites, while shrub cover and sagebrush cover were well below cover at the nest sites. Following conifer removal, we measured increases in herbaceous vegetation, primarily grasses, and sagebrush height. Our results indicate that conifer abundance can decrease habitat suitability for nesting sage‐grouse. Additionally, conifer removal can improve habitat suitability for nesting sage‐grouse within 3 years, and trajectories indicate that the habitat may continue to improve in the near future.     

Assessing lek attendance of male greater sage-grouse using fine-resolution GPS data: Implications for population monitoring of lek mating grouse

Counts of males displaying on breeding grounds are the primary management tool used to assess population trends in lekking grouse species. Despite the importance of male lek attendance (i.e., proportion of males on leks available for detection) influencing lek counts, patterns of within season and between season variability in attendance rates are not well understood. We used high-frequency global positioning system (GPS) telemetry data from male greater sage-grouse (Centrocercus urophasianus; n = 67) over five lekking seasons (2013–2017) at eight study sites in Nevada to estimate lek attendance rates. Specifically, we recorded daily locations of sage-grouse in relation to mapped lek boundaries and used generalized additive models to assess temporal variation in attendance rates by age class (subadult vs. adult). Average timing of peak attendance occurred on 16 April but varied from March 16, 2014 to April 21 , 2016. Overall, adult males attended leks at higher rates (0.683 at peak) and earlier in the season (19 March) than subadults (0.421 at peak on April 19). Peak attendance probability was positively related to cumulative winter precipitation. Daily probabilities of lek switching differed between adults (0.019 at peak on March 3) and subadults (0.046 at peak on March 22), and lek switching was negatively related to distance to nearest lek. Our results indicate variable patterns in lek attendance through time, and that lek switching may occur at higher rates than previously thought. We demonstrate the use of generalizable daily attendance curves to date-correct lek counts and derive estimates of male abundance, although such an approach will likely require the incorporation of information on age structure to produce robust results that are useful for population monitoring.     

Disturbance factors influencing greater sage-grouse lek abandonment in north-central Wyoming

Detecting the disappearance of active leks is the most efficient way to determine large declines in greater sage-grouse (Centrocercus urophasianus) populations; thus, understanding factors that influence lek abandonment is critical. We evaluated factors that may have influenced the probability of sage-grouse lek abandonment in the Bighorn Basin (BHB) of north-central Wyoming from 1980 to 2009. Our objective was to examine lek abandonment based on landscape characteristics that explain differences between occupied and unoccupied leks. We evaluated lek abandonment from 144 occupied and 39 unoccupied leks from the Wyoming Game and Fish Department lek database with sufficient data for our 30-year analysis. We conducted our analysis with binary logistic regression using landscape predictor variables obtained from geographic coverages at 5 scales (1.0-, 3.2-, 4.0-, 5.0-, and 6.4-km radii around leks) to evaluate how these disturbances have influenced lek abandonment. Coverages included anthropogenic characteristics such as agricultural development, oil and gas development, prescribed burned treatments, and roads; and environmental characteristics such as vegetation attributes and wildfire. Our combined model included the number of oil and gas wells in a 1.0-km radius, percent area of wildfire in a 1.0-km radius, and variability in shrub height in a 1.0-km radius around sage-grouse leks. Abandoned (unoccupied) leks had 1.1-times the variability of shrub height in a 1.0-km radius, 3.1-times the percentage of wildfire in a 1.0-km radius, and 10.3-times the number of oil and gas wells in a 1.0-km radius compared to occupied leks. The model-averaged odds of lek persistence with every 1 unit increase in oil and gas wells within a 1.0-km radius was 0.66 (90% CI: 0.37–0.94), odds with every 1% increase in wildfire in a 1.0-km radius was 0.99 (90% CI: 0.85–1.12), and odds with every 1 unit increase in the standard deviation of shrub height within a 1.0-km radius around a lek was 0.77 (90% CI: 0.45–1.08). Because the 90% confidence intervals around the odds ratios of wells did not overlap 1.0, we suggest this predictor variable was most influential in our model-averaged estimates. The BHB has lower developed reserves of oil and gas than many other regions; however, our study supports findings from other studies that demonstrate energy development increases lek abandonment. Our findings indicate conservation efforts should be focused on minimizing well development and implementing wildfire suppression tactics near active sage-grouse leks. © 2012 The Wildlife Society.     

Using gas chromatography to determine winter diets of greater sage-grouse in Utah

Sagebrush (Artemisia spp.) constitutes the majority (>99%) of sage-grouse (Centrocercus spp.) winter diets. Thus, identification and protection of important winter habitats is a conservation priority. However, not all sagebrush may be alike. More information is needed regarding sage-grouse sagebrush winter dietary preferences for application to management. The objective of our research was to determine if chemical analysis of fecal pellets could be used to characterize winter sage-grouse diets as a substitute for more invasive methods. We collected and analyzed fecal pellets and sagebrush samples from 29 different sage-grouse flock locations in northwestern and southcentral Utah. Using gas chromatography, we were able to identify crude terpene profiles that were unique to Wyoming sagebrush (A. tridentata wyomingensis) and black sagebrush (A. nova). We subsequently used the profiles to determine sagebrush composition of sage-grouse fecal pellets, thus reflecting sage-grouse winter diets. This technique provides managers with a tool to determine which species or subspecies of sagebrush may be important in the winter diets of sage-grouse populations. © 2011 The Wildlife Society.