Prey behavioral responses to predation risk in wolf‐ungulate‐plant systems are of interest to wildlife managers. Using Global Positioning System data collected from telemetry‐collared elk (Cervus elaphus) and wolves (Canis lupus), we evaluated elk behavioral responses to spatial and temporal variation in wolf‐ and human‐predation risk on a winter range in the Greater Yellowstone Area, USA. We found elk changed grouping patterns and increased movement rates as predation risk increased and that these behavioral changes were habitat dependent. Elk behavioral responses to wolf‐ and human‐predation risk were similar; however, responses to human‐predation risk were stronger than responses to wolf‐predation risk. These results suggest that predation risk from wolves or human hunters may result in elk spending more time on private rangelands away from public‐land winter ranges, which may exacerbate problems of landowner tolerance of elk on livestock pastures. However, increased movement and changing grouping patterns on winter ranges may also disperse elk grazing impacts and lessen elk impacts on any one area.

Summary1. Well-informed management of harvested species requires understanding how changing ecological conditions affect demography and population dynamics, information that is lacking for many species. We have limited understanding of the relative influence of carnivores, harvest, weather and forage availability on elk Cervus elaphus demography, despite the ecological and economic importance of this species. We assessed adult female survival, a key vital rate for population dynamics, from 2746 radio-collared elk in 45 populations across western North America that experience wide variation in carnivore assemblage, harvest, weather and habitat conditions. 2. Proportional hazard analysis revealed that 'baseline' (i.e. not related to human factors) mortality was higher with very high winter precipitation, particularly in populations sympatric with wolves Canis lupus. Mortality may increase via nutritional stress and heightened vulnerability to predation in snowy winters. Baseline mortality was unrelated to puma Puma concolor presence, forest cover or summer forage productivity. 3. Cause-specific mortality analyses showed that wolves and all carnivore species combined had additive effects on baseline elk mortality, but only reduced survival by <2%. When human factors were included, 'total' adult mortality was solely related to harvest; the influence of native carnivores was compensatory. Annual total mortality rates were lowest in populations sympatric with both pumas and wolves because managers reduced female harvest in areas with abundant or diverse carnivores. ‡Present address: Natural Resources Canada, 506 Burnside Road W, Victoria, BC V8Z 1M5, Canada *Correspondence author. E-mail: jedediah.brodie@gmail.com †Authorship alphabetical after B. Johnson.© 2013 This article is a US Government work and is in the public domain in the USA. 2013, 50, 295-305 doi: 10.1111/1365-2664.12044 4. Mortality from native carnivores peaked in late winter and early spring, while harvest-induced mortality peaked in autumn. The strong peak in harvest-induced mortality during the autumn hunting season decreased as the number of native carnivore species increased. 5. Synthesis and applications. Elevated baseline adult female elk mortality from wolves in years with high winter precipitation could affect elk abundance as winters across the western US become drier and wolves recolonize portions of the region. In the absence of human harvest, wolves had additive, although limited, effects on mortality. However, human harvest, and its apparent use by managers to offset predation, primarily controls overall variation in adult female mortality. Altering harvest quotas is thus a strong tool for offsetting impacts of carnivore recolonization and shifting weather patterns on elk across western North America. Journal of Applied Ecology

Wildlife managers need reliable methods to estimate large carnivore densities and population trends; yet large carnivores are elusive, difficult to detect, and occur at low densities making traditional approaches intractable. Recent advances in spatial capture‐recapture (SCR) models have provided new approaches for monitoring trends in wildlife abundance and these methods are particularly applicable to large carnivores. We applied SCR models in a Bayesian framework to estimate mountain lion densities in the Bitterroot Mountains of west central Montana. We incorporate an existing resource selection function (RSF) as a density covariate to account for heterogeneity in habitat use across the study area and include data collected from harvested lions. We identify individuals through DNA samples collected by (1) biopsy darting mountain lions detected in systematic surveys of the study area, (2) opportunistically collecting hair and scat samples, and (3) sampling all harvested mountain lions. We included 80 DNA samples collected from 62 individuals in the analysis. Including information on predicted habitat use as a covariate on the distribution of activity centers reduced the median estimated density by 44%, the standard deviation by 7%, and the width of 95% credible intervals by 10% as compared to standard SCR models. Within the two management units of interest, we estimated a median mountain lion density of 4.5 mountain lions/100 km2 (95% CI = 2.9, 7.7) and 5.2 mountain lions/100 km2 (95% CI = 3.4, 9.1). Including harvested individuals (dead recovery) did not create a significant bias in the detection process by introducing individuals that could not be detected after removal. However, the dead recovery component of the model did have a substantial effect on results by increasing sample size. The ability to account for heterogeneity in habitat use provides a useful extension to SCR models, and will enhance the ability of wildlife managers to reliably and economically estimate density of wildlife populations, particularly large carnivores.

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Traditional elk habitat management on public land has focused on providing security habitat for bull elk during the hunting season to provide for both adequate hunter opportunity and bull survival. This paradigm has given less consideration to adult female elk habitat use, patterns of adjacent land ownership, and hunter access. This paradigm also was developed when elk population sizes were much smaller in many areas. In many Rocky Mountain states, the focus of elk population management has recently shifted to reducing or maintaining elk population sizes, necessitating a better understanding of the implications of security habitat management, as well as patterns of adjacent land ownership and hunter access, on adult female elk. We addressed this need by testing the hypotheses that during the hunting season: 1) adult female elk selection for areas prohibiting or limiting hunter access is stronger than elk selection for publicly owned and managed elk security habitat, 2) these effects occur during the archery hunting period and intensify during the rifle hunting period, and 3) the effects of hunter access on selection are consistent among herds that occupy landscapes characterized by a matrix of public and private lands. We used global position system locations collected from 82 females in 2 different Greater Yellowstone Ecosystem (GYE) elk herds to evaluate effects of hunter access, security habitat as defined by the Hillis paradigm, and other landscape attributes on adult female elk resource selection during the pre‐hunting, archery, rifle, and post‐hunting periods. We found that female elk selection for areas restricting public hunting access was stronger than selection for security habitat in both study areas, and that the density of roads open to motorized use was the strongest predictor of elk distribution. Increases in selection for areas that restricted hunting access occurred during the rifle hunting period, and we did not find consistent evidence these movements were triggered by the archery hunting period. Our results provide evidence that in landscapes characterized by a matrix of public and privately owned lands, traditional concepts of elk security habitat need to be expanded to also include areas that restrict hunter access to plan for elk population management that is regulated through adult female harvest. Future efforts should investigate whether elk use of areas that restrict hunter access are flexible behavioral responses to hunting risk, or if these behaviors are passed from generation to generation such that a learned pattern of private land use becomes the normal movement pattern rather than a short‐term behavioral response. Published 2013. This article is a U.S. Government work and is in the public domain in the USA.

Anthrax, caused by the spore-forming bacterium Bacillus anthracis, is a zoonotic disease that affects humans and animals throughout the world. In North America, anthrax outbreaks occur in livestock and wildlife species. Vaccine administration in wildlife is untenable; the most effective form of management is surveillance and decontamination of carcasses. Successful management is critical because untreated carcasses can create infectious zones increasing risk for other susceptible hosts. We studied the bacterium in a re-emerging anthrax zone in southwest Montana. In 2008, a large anthraxepizootic primarily affected a domestic bison (Bison bison) herd and the male segment of a free-ranging elk (Cervus elaphus) herd in southwestern Montana. Following the outbreak, we initiated a telemetry study on elk to evaluate resource selection during the anthrax season to assist with anthrax management. We used a mixed effects generalized linear model (GLM) to estimate resource selection by male elk, and we mapped habitat preferences across the landscape. We overlaid preferred habitats on ecological niche model-based estimates of B. anthracis presence. We observed significant overlap between areas with a high predicted probability of male elk selection and B. anthracis potential. These potentially risky areas of elk and B. anthracis overlap were broadly spread over public and private lands. Future outbreaks in the region are probable, and this analysis identified the spatial extent of the risk area in the region, which can be used to prioritize anthrax surveillance.

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Fall elk (Cervus canadensis) habitat management on public lands provides security areas for reasonable elk survival and hunter opportunity. The management focus of maintaining or improving security areas, combined with conservative harvest regulations, may explain why some elk populations have increased in the western United States. However, in areas that include lands that restrict public hunter access, elk may alter their space use patterns during the hunting season by increasing use of areas that restrict public hunter access rather than using security areas on adjacent public lands. We used global positioning system location data from 325 adult female elk in 9 southwest Montana populations to determine resource selection during the archery and rifle hunting seasons. We found that during the archery season, in order of decreasing strength of selection, elk selected for areas that restricted access to public hunters, had greater time-integrated normalized difference vegetation index values, had higher canopy cover, were farther from motorized routes, and had lower hunter effort. During the rifle season, in order of decreasing strength of selection, elk selected for areas that restricted access to public hunters, were farther from motorized routes, had higher canopy cover, and had higher hunter effort. Interactions among several covariates revealed dependencies in elk resource selection patterns. Further, cross-population analyses revealed increased elk avoidance of motorized routes with increasing hunter effort during both the archery and rifle hunting seasons. We recommend managing for areas with !13% canopy cover that are !2,760 m from motorized routes, and identifying and managing for areas of high nutritional resources within these areas to create security areas on public lands during archery season. During the rifle season, we recommend managing for areas with !9% canopy cover that are !1,535 m from motorized routes, and are !20.23 km 2 . Lastly, given increased elk avoidance of motorized routes with higher hunter effort, we recommend that to maintain elk on public lands, managers consider increasing the amount of security in areas that receive high hunter effort, or hunting seasons that limit hunter effort in areas of high motorized route densities. Ó