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Shawn T. O’Neil

Publications and source records attributed to Shawn T. O’Neil.

35 records · Page 2Linked to original sources

A Bayesian multi-stage modelling framework to evaluate impacts of energy development on wildlife populations: An application to Greater Sage-Grouse (Centrocercus urophasianus)

Increased demand for domestic production of renewable energy has led to expansion of energy infrastructure across western North America. Much of the western U.S. comprises remote landscapes that are home to a variety of vegetation communities and wildlife species, including the imperiled sagebrush ecosystem and indicator species such as greater sage-grouse ( Centrocercus urophasianus ). Geothermal sources in particular have potential for continued development across the western U.S. but impacts to greater sage-grouse and other species are unknown. To address this information gap, we describe a novel two-pronged methodology that analyzes impacts of geothermal energy production on pattern and process of greater sage-grouse populations using (a) before-after control-impact (BACI) measures of population growth and lek absence rates and (b) concurrent-to-operation evaluations of demographic rates. Growth and absence rate analyses utilized 14 years of lek survey data collected prior (2005–2011) and concurrent (2012–2018) to geothermal operations at two sites in Nevada, USA. Demographic analyses utilized relocation data, restricted inference to concurrent years, and incorporated 17 additional control sites. Demographic results were applied to >100 potential geothermal sites distributed across the study region to generate spatially explicit predictions of unrealized population-level impacts.

MethodsX↗

Moisture abundance and proximity mediate seasonal use of mesic areas and survival of greater sage-grouse broods

Water is a critical and limited resource, particularly in the arid West, but water availability is projected to decline even while demand increases due to growing human populations and increases in duration and severity of drought. Mesic areas provide important water resources for numerous wildlife species, including the greater sage-grouse ( Centrocercus urophasianus ; hereafter, sage-grouse), an indicator for the health of sagebrush ecosystems. Understanding how wildlife use these crucial areas is necessary to inform management and conservation of sensitive species. Specifically, the influence of anthropogenic water subsidies such as irrigated pastures is not well-studied. We evaluated brood-rearing habitat selection and brood survival of sage-grouse in Long Valley, California, an area where the water rights are primarily owned by the city of Los Angeles and water is used locally to irrigate for livestock. This area thus represents a unique balance between the needs of wildlife and people that could increasingly define future water management. In this study, sage-grouse broods moved closer to the edge of mesic areas and used more interior areas during the late brood-rearing period, selecting for greener areas after 1 July. Mesic areas were particularly important during dry years, with broods using areas farther interior than in wet years. Brood survival was also positively influenced by the availability and condition of mesic resources, as indicated by variation in values of normalized difference vegetation index (NDVI), with survival peaking at moderate values of NDVI and just outside the edge but decreasing inside the mesic areas. Our results highlight the importance of quality edge habitat of large mesic areas for sage-grouse to balance habitat selection and survival, particularly during drier years and during the late brood-rearing period, which is a critical period because chick survival has been shown to influence population growth. This study highlights the implications of large-scale anthropogenic water manipulation, and the balance between local irrigation and water distribution to benefit other regions, from the context of a species of high conservation concern in North American sagebrush ecosystems.

California↗

Targeting sagebrush (Artemisia spp.) restoration following wildfire with Greater Sage-Grouse (Centrocercus urophasianus) nest selection and survival models

Unprecedented conservation efforts for sagebrush ( Artemisia spp.) ecosystems across the western United States have been catalyzed by risks from escalated wildfire activity that reduces habitat for sagebrush-obligate species such as Greater Sage-Grouse ( Centrocercus urophasianus ). However, post-fire restoration is challenged by spatial variation in ecosystem processes influencing resilience to disturbance and resistance to non-native invasive species, and spatial and temporal lags between slower sagebrush recovery processes and faster demographic responses of sage-grouse to loss of important habitat. Decision-support frameworks that account for these factors can help users strategically apply restoration efforts by predicting short and long-term ecological benefits of actions. Here, we developed a framework that strategically targets burned areas for restoration actions (e.g., seeding or planting sagebrush) that have the greatest potential to positively benefit sage-grouse populations through time. Specifically, we estimated sagebrush recovery following wildfire and risk of non-native annual grass invasion under four scenarios: passive recovery, grazing exclusion, active restoration with seeding, and active restoration with seedling transplants. We then applied spatial predictions of integrated nest site selection and survival models before wildfire, immediately following wildfire, and at 30 and 50 years post-wildfire based on each restoration scenario and measured changes in habitat. Application of this framework coupled with strategic planting designs aimed at developing patches of nesting habitat may help increase operational resilience for fire-impacted sagebrush ecosystems.

Environmental Management↗

Invasion of annual grasses following wildfire corresponds to maladaptive habitat selection by a sagebrush ecosystem indicator species

Numerous wildlife species within semi-arid shrubland ecosystems across western North America are experiencing substantial habitat loss and fragmentation. These changes in habitat are often attributed to a diverse suite of factors including prolonged and increasingly severe droughts, conifer expansion, anthropogenic development, domestic and feral livestock grazing, and invasion of exotic annual grasses, which promotes increased wildfire frequency and severity. Greater sage-grouse ( Centrocercus urophasianus ; hereafter, sage-grouse) are considered an indicator of sagebrush ecosystem health and have experienced widespread population decline associated with habitat loss and degradation, as well as changes in predator communities. Our objectives were to model and map sage-grouse habitat selection and survival during the important brood-rearing life stage in relation to landscape-scale environmental predictors. Furthermore, we sought to understand impacts of wildfire and annual grass invasion on brood habitat, as these accelerated disturbance regimes are a primary cause of habitat loss within the Great Basin region of the USA. We used a hierarchical Bayesian modeling framework to estimate resource selection functions and survival for early and late brood-rearing stages of sage-grouse in relation to a broad suite of habitat characteristics evaluated at multiple spatial scales within the Great Basin from 2009 to 2019. Sage-grouse selected for greater perennial grass cover, higher relative elevations, and areas closer to springs and wet meadows during both early and late brood-rearing. Terrain characteristics, including heat load and aspect, were important in survival models, as was variation in shrub height. We also found strong evidence for higher survival for both early and late broods within previously burned areas, but survival within burned areas decreased as annual grass cover (i.e. cheatgrass , Bromus tectorum ) increased. This interaction effect demonstrates how invasion of annual grasses into burned areas, which has become prevalent in Great Basin sagebrush ecosystems, can lead to maladaptive habitat selection by brood-rearing greater sage-grouse. Understanding these complex relationships aids wildlife conservation and habitat management as wildfire and annual grass cycles continue to accelerate across western ecosystems.

California, Nevada↗

Behavioral state-dependent habitat selection and implications for animal translocations

Post-release monitoring of translocated animals is often used to inform future translocation protocols. Quantifying habitat selection of translocated individuals may help identify features that characterize good settlement habitat and thus inform the choice of future release sites. However, translocated animals often undergo post-release behavioural modification, and their habitat selection may vary depending on the underlying behavioural state. To investigate this, we analysed behavioural state-dependent habitat selection in female greater sage-grouse Centrocercus urophasianus translocated from Wyoming to North Dakota, USA, using Hidden Markov Models combined with Integrated Step Selection Analysis. We segmented individual trajectories into behavioural phases corresponding to an exploratory state, characterized by broad and directed movements, and a restricted state, characterized by short and tortuous movements. Then, we quantified habitat selection in each state while accounting for seasonality and individual reproductive status. While in the exploratory state, sage-grouse exhibited natal habitat preference induction by selecting for high sagebrush cover, which is typical of their natal area in Wyoming but not of the release area in North Dakota. In the restricted state, sage-grouse selected for gentle topography and also adjusted their habitat selection to constraints imposed by seasonality and reproductive needs by selecting for high herbaceous cover during brood rearing. Synthesis and applications . Habitat selection of translocated sage-grouse differed between the post-release exploration and the settlement phase. Features selected after settling, not during exploration, are likely indicative of suitable settlement habitat. Our results suggest that areas characterized by gentle topography and high herbaceous cover are well-suited as release sites for sage-grouse translocated during the summer, especially brood-rearing females, and that sagebrush cover may not be a critical factor in determining the appropriateness of release sites for sage-grouse in North Dakota. Our findings highlight the need to consider behaviour when using habitat selection estimates to inform the choice of future release sites.

North Dakota, Wyoming↗

Sage-grouse population dynamics are adversely impacted by overabundant feral horses

In recent decades, feral horse ( Equus caballus ; horse) populations increased in sagebrush ( Artimesia spp.) ecosystems, especially within the Great Basin, to the point of exceeding maximum appropriate management levels (AML max ), which were set by land administrators to balance resource use by feral horses, livestock, and wildlife. Concomitantly, greater sage-grouse ( Centrocercus urophasianus ; sage-grouse) are sagebrush obligates that have experienced population declines within these same arid environments as a result of steady and continued loss of seasonal habitats. Although a strong body of research indicates that overabundant populations of horses degrade sagebrush ecosystems, empirical evidence linking horse abundance to sage-grouse population dynamics is missing. Within a Bayesian framework, we employed state-space models to estimate population rate of change ( λ ) using 15 years (2005–2019) of count surveys of male sage-grouse at traditional breeding grounds (i.e., leks) as a function of horse abundance relative to AML max and other environmental covariates (e.g., wildfire, precipitation, % sagebrush cover). Additionally, we employed a post hoc impact-control design to validate existing AML max values as related to sage-grouse population responses, and to help control for environmental stochasticity and broad-scale oscillations in sage-grouse abundance. On average, for every 50% increase in horse abundance over AML max , our model predicted an annual decline in sage-grouse abundance by 2.6%. Horse abundance at or below AML max coincided with sage-grouse λ estimates that were consistent with trends at non-horse areas elsewhere in the study region. Thus, AML max , as a whole, appeared to be set adequately in preventing adverse effects to sage-grouse populations. Results indicated 76%, 97%, and >99% probability of sage-grouse population decline relative to controls when horse numbers are 2, 2.5, and ≥3 times over AML max , respectively. As of 2019, horse herds exceeded AML max in Nevada, USA, by >4 times on average across all horse management areas. If feral horse populations continue to grow at current rates unabated, model projections indicate sage-grouse populations will be reduced within horse-occupied areas by >70.0% by 2034 (15-year projection), on average compared to 21.2% estimated for control sites. A monitoring framework that improves on estimating horse abundance and identifying responses of sage-grouse and other key indicator species (plant and animal) would be beneficial to guide management decisions that promote co-occurrence of horses with sensitive wildlife and livestock within landscapes subjected to multiple uses. Published 2021. This article is a U.S. Government work and is in the public domain in the USA. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.

Nevada↗

Large-scale wildfire reduces population growth in a peripheral population of sage-grouse

Drastic increases in wildfire size and frequency threaten western North American sagebrush ( Artemisia L. spp.) ecosystems. At relatively large spatial scales, wildfire facilitates type conversion of sagebrush-dominated plant communities to monocultures of invasive annual grasses ( e.g., Bromus tectorum L.). Annual grasses provide fine fuels that promote fire spread, contributing to a positive grass–fire feedback cycle that affects most sagebrush ecosystems, with expected habitat loss for resident wildlife populations. Greater sage-grouse ( Centrocercus urophasianus Bonaparte, 1827) are sagebrush obligate species that are indicators of sagebrush ecosystem function because they rely on different components of sagebrush ecosystems to meet seasonal life history needs. Because wildfire cannot be predicted, chronic impacts of wildfire on sage-grouse populations have been largely limited to correlative studies. Thus, evidence from well-designed experiments is needed to understand the specific mechanisms by which wildfire is detrimental to sage-grouse population dynamics.

California, Nevada↗

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.

Montana, North Dakota, South Dakota, Wyoming↗

A rapid assessment function to estimate common raven population densities: Implications for targeted management

Common raven ( Corvus corax ; raven) populations have increased over the past 5 decades within the western United States. Raven population increases have been largely attributed to growing resource subsidies from expansion of human enterprise. Concomitantly, managers are becoming increasingly concerned about elevated adverse effects on multiple sensitive prey species, damage to livestock and agriculture, and human safety. Managers could benefit from a rapid but reliable method to estimate raven densities across spatiotemporal scales to monitor raven populations more efficiently and inform targeted and adaptive management frameworks. However, obtaining estimates of raven density is data- and resource-intensive, which renders monitoring within an adaptive framework unrealistic. To address this need, we developed a rapid survey protocol for resource managers to estimate site-level density based on the average number of ravens per survey. Specifically, we first estimated raven densities at numerous field sites with robust distance sampling procedures and then used regression to investigate the relationship between those density estimates and the number of ravens per survey, which revealed a strong correlation ( R 2 = 0.86). For management application, we provide access to R function software through a web-based interface to estimate density using number of ravens per survey, which we refer to as a Rapid Assessment Function (RAF). Then, using a simulation analysis of data from sites with abundant surveys and the RAF, we estimated raven density based on different numbers of surveys to help inform how many surveys are needed to achieve reliable estimates within this rapid assessment. While more robust procedures of distance sampling are the preferred methods for estimating raven densities from count surveys, the RAF tool presented herein provides a reliable approximation for informing management decisions when managers are faced with resource and small sample size constraints.

California, Idaho, Nevada, Oregon↗

SMaRT: A science-based tiered framework for common ravens

Large-scale increases and expansion of common raven ( Corvus corax ; raven) populations are occurring across much of North America, leading to increased negative consequences for livestock and agriculture, human health and safety, and sensitive species conservation. We describe a science-based adaptive management framework that incorporates recent quantitative analyses and mapping products for addressing areas with elevated raven numbers and minimizing potential adverse impacts to sensitive species, agricultural damage, and human safety. The framework comprises 5 steps: (1) desktop analysis; (2) field assessments; (3) comparison of raven density estimates to an ecological threshold (in terms of either density or density plus distance to nearest active or previous nest); (4) prescribing management options using a 3-tiered process (i.e., habitat improvements, subsidy reductions, and direct actions using StallPOPd.V4 software); and (5) post-management monitoring. The framework is integrated within the Science-based Management of Ravens Tool (SMaRT), a web-based application outfitted with a user-friendly interface that guides managers through each step to develop a fully customized adaptive plan for raven management. In the SMaRT interface, users can: (1) interact with pre-loaded maps of raven occurrence and density and define their own areas of interest within the Great Basin to delineate proposed survey or treatment sites; (2) enter site-level density estimates from distance sampling methods or perform estimation of raven densities using the rapid assessment protocol that we provide; (3) compare site-level density estimates to an identified ecological threshold; and (4) produce a list of potential management options for their consideration. The SMaRT supports decision-making by operationalizing scientific products for raven management and facilitates realization of diverse management goals including sensitive species conservation, protection of livestock and agriculture, safeguarding human health, and addressing raven overabundance and expansion. We illustrate the use of the framework through SMaRT using an example of greater sage-grouse ( Centrocercus urophasianus ) conservation efforts within the Great Basin, USA.

California, Idaho, Nevada, Oregon, Utah↗

Spatial modeling of common raven density and occurrence helps guide landscape management within Great Basin sagebrush ecosystems

Common ravens ( Corvus corax ; ravens) are a behaviorally flexible nest predator of several avian species, including species of conservation concern. Movement patterns based on life history phases, particularly territoriality of breeding birds and transiency of nonbreeding birds, are thought to influence the frequency and efficacy of nest predation. As such, predicting where on the landscape territorial resident and non-territorial transient birds may be found in relation to the distribution of sensitive prey is of increasing importance to managers and conservationists. From 2007 to 2019, we conducted raven point count surveys between mid-March and mid-September across 43 different field sites representing typical sagebrush ( Artemisia spp.) ecosystems of the Great Basin, USA. The surveys conducted during 2007–2016 were used in previously published maps of raven occurrence and density. Here, we examined the relationship between occurrence and density of ravens using spatially explicit predictions from 2 previously published studies and differentiate areas occupied by higher concentrations of resident ravens as opposed to transients. Surveys conducted during 2017–2019 were subsequently used to evaluate the predicted trends from our analytical approach. Specifically, we used residuals from a generalized linear regression to establish the relationship between occurrence and density, which ultimately resulted in a spatially explicit categorical map that identifies areas of resident versus transient ravens. We evaluated mapped categories using independently collected observed raven group sizes from the 2017–2019 survey data, as well as an independent dataset of global positioning system locations of resident and transient individuals monitored during 2019–2020. We observed moderate agreement between the mapped categories and independent datasets for both evaluation approaches. Our map provides broad inference about spatial variation in potential predation risk from ravens for species such as greater sage-grouse ( Centrocercus urophasianus ) and can be used as a valuable spatial layer for decision support tools aimed at guiding raven management decisions and, ultimately, improving survival and reproduction of sensitive prey within the Great Basin.

Arizona, California, Idaho, Montana, Nevada, Orego↗

Inter- and intra-annual effects of lethal removal on common raven abundance in Nevada and California, USA

Populations of common ravens ( Corvus corax ; ravens) have increased rapidly within sagebrush ( Artemisia spp.) ecosystems between 1960 and 2020. Although ravens are native to North America, their population densities have expanded to levels that negatively influence the population dynamics of other wildlife species of conservation concern, such as greater sage-grouse ( Centrocercus urophasianus ) and desert tortoises ( Gopherus agassizii ). For this reason, lethal removal, such as the application of the avicide DRC-1339, has been used to manage raven numbers at local scales and under certain circumstances. Because the relative effectiveness of DRC-1339 in reducing raven populations densities is not thoroughly understood, we completed 2 case studies using a before-after-control-impact experimental design of density estimates generated from point count data within a Bayesian hierarchical distance sampling framework. Specifically, we analyzed >16,000 point count surveys collected during 2009–2019 and split into 2 study designs covering multiple field sites within the Great Basin region. The first experiment evaluated intra-annual changes in density by comparing before and after treatment time periods within a single breeding season for multiple treatment regions compared to 2 control regions. The other experiment focused on inter-annual differences by comparing time periods across years before and after the onset of annual avicide application for a single treatment region compared to multiple control regions. Our models estimated a 100% probability of decline in density relative to control sites for both the intra- and inter-annual model designs. At treatment sites, expected densities of ravens varied but were reduced by 43% (95% CRI: 33–49%) and 54% (95% CRI: 24–71%) according to intra- and inter-annual analyses, respectively, whereas densities increased by 42% (95% CRI: 27–60%) and 15% (95% CRI: -17 to 58%) at control sites. Although population densities were reduced with treatments, trends indicated that sustained effort would likely be needed to maintain densities at acceptable levels within regions of interest. Effectively reducing the adverse effects of raven populations on other native species likely will depend on a variety of targeted management actions such as improving habitat quality for prey species, possibly reducing ravens’ population density, and treating the cause of increased raven abundance to reduce future carrying capacity and prevent rebounds.

California, Nevada↗

Wildfire and the ecological niche: Diminishing habitat suitability for an indicator species within semi‐arid ecosystems

Globally accelerating frequency and extent of wildfire threatens the persistence of specialist wildlife species through direct loss of habitat and indirect facilitation of exotic invasive species. Habitat specialists may be especially prone to rapidly changing environmental conditions because their ability to adapt lags behind the rate of habitat alteration. As a result, these populations may become increasingly susceptible to ecological traps by returning to suboptimal breeding habitats that were dramatically altered by disturbance. We demonstrate a multistage modeling approach that integrates habitat selection and survival during the key nesting life‐stage of a bird species of high conservation concern, the greater sage‐grouse ( Centrocercus urophasianus ; hereafter, sage‐grouse). We applied these spatially explicit models to a spatiotemporally robust dataset of sage‐grouse nest locations and fates across wildfire‐altered sagebrush ecosystems of the Great Basin ecoregion, western United States. Female sage‐grouse exhibited intricate habitat selection patterns that varied across regional gradients of ecological productivity among sagebrush communities, but often selected nest sites that disproportionately resulted in nest failure. For example, 23% of nests occurred in wildfire‐affected habitats characterized by reduced sagebrush cover and greater composition of invasive annual grasses. We found survival of nests was negatively associated with wildfire‐affected areas, but positively associated with higher elevations with increased ruggedness and overall shrub cover. Strong site fidelity likely drove sage‐grouse to continue nesting in habitats degraded by wildfire. Hence, increasing frequency and extent of wildfire may contribute disproportionately to reduced reproductive success by creating ecological traps that act as population sinks. Identifying such habitat mismatches between selection and survival facilitates deeper understanding of the mechanisms driving reduced geographic niche space and population decline at broad spatiotemporal scales, while guiding management actions to areas that would be most beneficial to the species.

Global Change Biology↗

Population and habitat analyses for greater sage-grouse (Centrocercus urophasianus) in the bi-state distinct population segment—2018 update

Executive Summary The Bi-State Distinct Population Segment (Bi-State DPS) of greater sage-grouse ( Centrocercus urophasianus , hereinafter “sage-grouse”) represents a genetically distinct and geographically isolated population that straddles the border between Nevada and California. The primary threat to these sage-grouse populations is the expansion of single-leaf pinyon ( Pinus monophylla ) and Utah juniper ( Juniperus osteosperma ) into sagebrush ecosystems, which fragments and reduces population connectivity and survival. Other important threats include low water availability during brood-rearing, particularly during drought, and increased predation by common ravens ( Corvus corax ), a generalist predator often associated with anthropogenic resource subsidies. Although the Bi-State DPS occurs at high elevations relative to sage-grouse range-wide, changes in historical wildfire cycles and the conversion of native shrubs to invasive annual grasslands still threaten these populations. The Bi-State DPS has undergone multiple federal status assessments and associated litigation. For example, in October of 2013, the Bi-State DPS was proposed for listing as threatened under the Endangered Species Act of 1973 by the U.S. Fish and Wildlife Service (USFWS), then withdrawn in April 2015. The withdrawal decision was challenged, and in May 2018, a Federal district court ordered the withdrawal decision to be vacated, and USFWS was required to re-open the October 2013 listing evaluation. In response, the U.S. Geological Survey (USGS), with State and Federal collaborators, embarked on a multipronged analysis to provide current and best available science regarding population status of sage-grouse within the Bi-State DPS. Using data from a long-term monitoring program, we carried out four analytical study objectives, and here, we provide preliminary results of these analyses. First, we used integrated population modeling (IPM) to predict annual population abundance and annual finite rate of population change for the Bi-State DPS, as a whole, and for each subpopulation between 1995 and 2018. Because sage-grouse exhibit population cycles (periodic increases and decreases in abundance across approximately 6- to 10-year wavelengths), we estimated trends across three nested temporal scales that represent one (11 years), two (18 years), and three (24 years) complete population cycles. These estimates of relatively long-term averaged population change account for temporal (that is, interannual) variation. Our model predicted population abundance for the Bi-State DPS during 2018 at 3,305 individuals (2,247–4,683), with the majority occupying Bodie Hills and Long Valley. The model also predicted cyclic dynamics in abundance through time with evidence of 24-year population growth and slight trends of decline over the past 18 years. Specifically, across the Bi-State DPS as a whole, we estimated annual average at 0.99, 0.99, and 1.02 over the one, two, and three population cycles, which equated to a 10.5 percent, 16.6 percent decrease, and 60.0 percent increase in abundance over the 11-, 18-, and 24-year cycles. Estimated abundance in 2018 had not reached numbers lower than those predicted during 1995. However, we found spatial variation in population trends across the three cycles. Bodie Hills subpopulation comprised the greatest (1,521) and exhibited average annual greater than 1.0 across all periods resulting in average annual increases of 7 percent. This relatively large subpopulation has grown 5 times larger than what was predicted in 1995 while experiencing cyclical dynamics within that period. Conversely, other smaller subpopulations within the Bi-State DPS exhibited average annual equal to or less than 1.0 resulting in estimated 10-year risks of extirpation ranging from 2.0 to 76.1 percent. In general, evidence of decline among smaller subpopulations was greatest for the most recent period (2008–18) compared to a period that encompassed three full population cycles (24-year). This difference coincides with an intense period of drought that began in 2012. For comparative purposes as part of this first objective, we conducted a similar analysis for populations of sage-grouse within Nevada and California but outside the Bi-State DPS. We developed a region-wide and distance-weighted IPM using lek count from Nevada Department of Wildlife (NDOW) and California Department of Fish and Wildlife (CDFW) databases and with telemetry data collected by USGS across 12 sage-grouse subpopulations. Our models predicted similar patterns in population cycling outside the Bi-State DPS but with much stronger evidence of long-term declines across 24 years. Specifically, median averaged across each year of the 11-, 18-, and 24-year periods resulted in average annual values of 0.94, 0.97, and 0.99, respectively. These values equate to 41.0 percent, 38.5 percent, and 21.3 percent declines over the corresponding periods. Second, we used lek count data in a state-space modeling framework to compare trends in population abundance across different spatial scales (that is, leks versus Bi-State DPS). This hierarchical framework allowed us to disentangle declines associated with climate conditions as opposed to other local level factors that might signal the need for management intervention. Specifically, we identified 7 leks that were both declining and recently decoupled from larger spatial scale trends, typically governed by climatic conditions (referred to as soft or hard signals). The goal of this analysis was to provide an early warning system that might have implications for conservation actions at local scales. Third, we developed phenological (spring, summer–fall, and winter) and reproductive life stage (nesting, early brood-rearing, and late-brood rearing) based resource selection functions using various environmental covariates. We report rankings of variable importance for each season and life stage, developed maps of habitat selection indices (HSI), binned categories representing low, moderate, and high classes of quality (where any category greater than or equal to low indicated selected habitat) for each phenological season and life stage, and produced composite maps by selected phenological and reproductive stage to estimate annual habitat. Fourth, we used for each lek within the Bi-State DPS to carry out a spatial analysis that quantified substantial changes in the distribution of occupied habitat across long- (24-year) and short- (11-year) term periods. Owing to differences among available datasets, the long-term analysis primarily reflected spatial shifts among subpopulations comprising the majority of the Bi-State DPS (that is, Bodie Hills and Long Valley) while the short-term analysis also quantified changes among subpopulations along the periphery. Over long and short-term periods, the overall distribution of occupied habitat (as measured by 99 percent utilization distributions intersecting any quantified habitat) was reduced by 20,573 ha and 55,492 ha, respectively. Occupied core areas (as measured by 50 percent utilization distributions intersecting any quantified habitat) over long-term periods were solely located in Bodie Hills and Long Valley. Although nearly all subpopulations experienced contractions in occupied overall and core distribution, Bodie Hills experienced spatial expansion that occurred with concomitant spatial contraction at Long Valley over both periods. Subpopulations at the northern (Pine Nuts), central (Sagehen) and southern (White-Mountains) extents of the Bi-State DPS also experienced spatial contraction over the short-term period. These findings, coupled with those of population trends, indicate long-term patterns in redistribution of sage-grouse from Long Valley and peripheral subpopulations to Bodie Hills. That is, sage-grouse subpopulations at the periphery are declining while the largest population at the core is increasing, which could have meaningful impacts on overall metapopulation persistence. We provide evidence for loss of occupied habitat (reduced distribution) given local extirpation of subpopulations. Fifth, we calculated percentages of selected phenological, life stage, and annual habitat that each subpopulation contributed to the Bi-State DPS. We then intersected these maps with a composite estimate of occupied habitat from the fourth objective and calculated percentages of selected habitat likely occupied by sage-grouse that each subpopulation contributed to the Bi-State DPS. These values provide evidence for loss of occupied habitat and subsequent reductions in spatial distribution given reductions in abundance and, in some cases, extirpation of leks within subpopulations. Lastly, we carried out an initial in-depth analysis of selection for irrigated pastures and wet meadows during the brood-rearing stage for the Long Valley subpopulation. We chose this subpopulation because it represents a population core, representing 26.5 percent of total sage-grouse within the Bi-State DPS, and has exhibited long-term declines in abundance and distribution. This subpopulation is highly sensitive to precipitation and other factors that influence water availability. Models predicted higher use of the interior portions of irrigated pastures and wet meadows during late brood-rearing period, which represented a potentially risky use of habitat that was exacerbated during periods of low moisture (for example, drought, reduced water delivery, or both). Sage-grouse typically used edges of riparian areas and pastures, largely because the interior of these mesic areas consisted of considerably less overhead concealment cover (for example, shrubs) that likely resulted in a higher risk of mortality. We found that a lack of water delivery to pastures in the form of overwinter precipitation or diversion ditches increased the movements of sage-grouse to the interior of pastures. Although further investigation of water delivery impacts on chick survival are warrented, our initial findings regarding resource selection may explain recent declines in population growth at Long Valley.

California, Nevada↗

Broad‐scale occurrence of a subsidized avian predator: reducing impacts of ravens on sage‐grouse and other sensitive prey

Expanding human enterprise across remote environments impacts numerous wildlife species. Anthropogenic resources provide subsidies for generalist predators that can lead to cascading effects on prey species at lower trophic levels. A fundamental challenge for applied ecologists is to disentangle natural and anthropogenic influences on species occurrence, and subsequently develop spatially explicit models to help inform management and conservation decisions. Using Bayesian hierarchical occupancy models, we mapped the broad‐scale occurrence of common ravens Corvus corax as a function of natural and anthropogenic landscape covariates using >15,000 point count surveys performed during 2007–2016 within the Great Basin region, USA. Raven abundance and distribution is substantially increasing across the American west due to unintended anthropogenic resource subsidies. Importantly, ravens prey on eggs and chicks of numerous species including greater sage‐grouse Centrocercus urophasianus , an indicator species whose decline is at the centre of national conservation strategies and land use policies.Anthropogenic factors that contributed to greater raven occurrence were: increased road density, presence of transmission lines, agricultural activity, and presence of roadside rest areas. Natural landscape characteristics included lower elevations with greener vegetation (NDVI), greater stream and habitat edge densities, and lower percentages of big sagebrush A. tridentata spp . Interactions between anthropogenic sources of nesting substrate and food subsidies suggested that raven occurrence increased multiplicatively when these resource subsidies co‐occurred. Overall, the average probability of raven occurrence estimated within sagebrush ecosystems of the study area was ~0.83. Synthesis and applications . We demonstrate how anthropogenic factors can be disentangled from natural effects when making spatially‐explicit predictions of subsidized predators occurring across expansive landscapes. This approach can guide management decisions where subsidized predators overlap sensitive prey habitats. For example, we identify areas where elevated raven occurrence coincides with breeding sage‐grouse concentration areas and appears to be largely driven by anthropogenic factors. Management applications could focus on reducing raven access to anthropogenic subsidies in these areas, while prioritizing habitat improvements for sage‐grouse elsewhere. Our approach is applicable to other species where widespread survey data are available.

Journal of Applied Ecology↗

Habitat selection by postbreeding female diving ducks: Influence of habitat attributes and conspecifics

Habitat selection studies of postbreeding waterfowl have rarely focused on within-wetland attributes such as water depth, escape cover, and food availability. Flightless waterfowl must balance habitat selection between avoiding predation risks and feeding. Reproductively successful female ducks face the greatest challenges because they begin the definitive prebasic molt at or near the end of brood rearing, when their body condition is at a low point. We assessed the relative importance of habitat attributes and group effects in habitat selection by postbreeding female lesser scaup Aythya affinis on a 2332-ha montane wetland complex during the peak flightless period (August) over seven years. Hypothesis-based habitat attributes included percent open water, open water:emergent edge density, water depth, percent flooded bare substrate, fetch (distance wind can travel unobstructed), group size, and several interactions representing functional responses to interannual variation in water levels. Surveys of uniquely marked females were conducted within randomly ordered survey blocks. We fitted two-part generalized linear mixed-effects models to counts of marked females within survey blocks, which allowed us to relate habitat attributes to relative probability of occurrence and, given the presence of a marked female, abundance of marked individuals. Postbreeding female scaup selected areas with water depths > 40 cm, large open areas, and intermediate edge densities but showed no relation to flooded bare substrate, suggesting their habitat preferences were more influenced by avoiding predation risks and disturbances than in meeting foraging needs. Grouping behavior by postbreeding scaup suggests habitat selection is influenced in part by behavioral components and/or social information, conferring energetic and survival benefits (predation and disturbance risks) but potentially also contributing to competition for food resources. This study demonstrates the importance of incorporating group effects and interannual variability in habitat conditions when investigating habitat selection, particularly for seasons when waterfowl are aggregated.

Journal of Avian Biology↗

Behavioural cues surpass habitat factors in explaining prebreeding resource selection by a migratory diving duck

Prebreeding habitat selection in birds can often be explained in part by habitat characteristics. However, females may also select habitats on the basis of fidelity to areas of previous reproductive success or use by conspecifics. The relative influences of sociobehavioural attributes versus habitat characteristics in habitat selection has been primarily investigated in songbirds, while less is known about how these factors affect habitat selection processes in migratory waterfowl. Animal resource selection models often exhibit much unexplained variation; spatial patterns driven by social and behavioural characteristics may account for some of this. We radiomarked female lesser scaup, Aythya affinis , in the southwestern extent of their breeding range to explore hypotheses regarding relative roles of habitat quality, site fidelity and conspecific density in prebreeding habitat selection. We used linear mixed-effects models to relate intensity of use within female home ranges to habitat features, distance to areas of reproductive success during the previous breeding season and conspecific density. Home range habitats included shallow water (≤118 cm), moderate to high densities of flooded emergent vegetation/open water edge and open water areas with submerged aquatic vegetation. Compared with habitat features, conspecific female density and proximity to successful nesting habitats from the previous breeding season had greater influences on habitat use within home ranges. Fidelity and conspecific attraction are behavioural characteristics in some waterfowl species that may exert a greater influence than habitat features in influencing prebreeding space use and habitat selection within home ranges, particularly where quality habitat is abundant. These processes may be of critical importance to a better understanding of habitat selection in breeding birds.

Montana↗