Bark Beetle-Affected Forests Provide Elk Only a Marginal Refuge from Hunters

For nearly 2 decades, the forests of the Rocky Mountains in the United States experienced a bark-beetle (Dendoctronus ponderosae) epidemic. The number of dead and falling trees from this epidemic likely will affect how elk (Cervus canadensis) and hunters use the forest and their interactions. Downed trees potentially create a component of refuge habitat that could affect the effectiveness of hunting to regulate abundance of growing elk populations. We evaluated how forests affected by bark beetles in south-central Wyoming, USA, influenced resource selection of 50 female elk and 374 hunters in 2012–2016, and interactions between elk and hunters. We employed global positioning system (GPS) technology on elk and hunters, and developed a satellite-derived land classification specifically depicting beetle-affected forests. We tested the predictions that elk would increase use of beetle-killed areas from summer to the hunting season, and that hunters would avoid beetle-killed areas regardless of elk use. Elk increased use of beetle-killed areas during hunting seasons as did hunters during the archery season; however, during the rifle season, hunters avoided beetle-killed areas. Nevertheless, during the rifle season, areas of beetle-kill with a high probability of elk occurrence dampened the aversion hunters had towards beetle-killed areas. Therefore, in contrast to our expectations, forests that have been altered by the bark-beetle epidemic may only function marginally as a refuge for elk. Our study area was at the beginning of the tree-fall phase (i.e., ~3–7 yr after peak infestation) of the bark-beetle epidemic; thus, future research efforts should focus on how a continued increase in downed trees will influence interactions between elk and hunters. At current levels of tree fall, however, beetle-kill should not limit the ability of managers to regulate elk herds through harvest. © 2020 The Wildlife Society.     

Autumn resource selection by female elk in a recently burned landscape in western Montana

Wildfire activity across the western United States has increased in recent decades, with wildfires burning at a higher severity and larger scale. The effect of wildfires on forest structure and wildlife habitat is largely influenced by wildfire severity; however, few studies have evaluated the effects of wildfire severity on resource selection of ungulates, particularly during hunting seasons, when knowledge of resource selection is essential for making informed management decisions. To fill this knowledge gap, we fit resource selection probability functions for female elk (Cervus canadensis) in years 2 and 3 post-wildfire to evaluate the effects of wildfire severity and other environmental and anthropogenic factors on elk resource selection during 4 autumn periods with varying levels of hunter pressure (prehunt, archery-only, backcountry rifle, and rifle). The probability of female elk selecting low-severity burned forests during the prehunt, archery-only, backcountry rifle, and rifle periods was 0.99 (95% credible interval [CrI] = 0.98–1.00), 0.99 (CrI = 0.97–1.00), 0.99 (CrI = 0.99–1.00), and 0.0010 (CrI = 0.00067–0.0015]), respectively, and did not strongly differ from the probability of selecting high-severity burned forests. During the prehunt period, elk also selected areas with greater forage quality and areas farther from open roads. Elk selected similar resources during the archery period, and selected areas with higher hunter pressure. Elk started leaving hunting districts that had higher snowpack (i.e., snow water equivalent; β = −0.84, CrI = −0.96–−0.72) and allowed rifle hunting (β = −5.39, CrI = −5.80–−4.97) but still selected areas with higher hunter pressure (β = 0.92, CrI = 0.78–1.07) during the backcountry rifle period. During the rifle period, elk continued avoiding areas with high snowpack (β = −3.96, CrI = −4.22–−3.71) and started selecting areas with lower hunter pressure (β = −1.71, CrI = −1.79–−1.64) and lower canopy cover. Overall, wildfire affected elk distributions in early autumn 2 and 3 years after fire in our study area, with limited differences in resource selection between wildfire severity categories. By late autumn, hunter pressure and snowpack were the primary factors influencing elk distribution, and wildfire had little influence on selection. When estimating wildfire effects on elk movements during autumn and establishing appropriate hunting regulations, managers should consider the hunting season, hunter pressure, timing and amount of snowpack, location of traditional winter range, and the seasonal elk range burned, as all these factors may contribute to how elk use the landscape in autumn.     

Sex-Specific Elk Resource Selection during the Anthrax Risk Period

Anthrax, caused by the spore-forming bacterium Bacillus anthracis, is a zoonosis affecting animals and humans globally. In the United States, anthrax outbreaks occur in wildlife and livestock, with frequent outbreaks in native and exotic wildlife species in Texas, livestock outbreaks in the Dakotas, and sporadic mixed outbreaks in Montana. Understanding where pathogen and host habitat selection overlap is essential for anthrax management. Resource selection and habitat use of ungulates may be sex-specific and lead to differential anthrax exposure risks across the landscape for males and females. We evaluated female elk (Cervus canadensis) resource selection in the same study areas as male elk in a previous anthrax risk study to identify risk of anthrax transmission to females and compare transmission risk between females and males. We developed a generalized linear mixed-effect model to estimate resource selection for female elk in southwest Montana during the June to August anthrax transmission risk period. We then predicted habitat selection of female and male elk across the study area and compared selection with the distribution of anthrax risk to identify spatial distributions of potential anthrax exposure for the male and female elk. Female and male elk selected different resources during the anthrax risk period, which resulted in different anthrax exposure areas for females and males. The sex-specific resource selection and habitat use could infer different areas of risk for anthrax transmission, which can improve anthrax and wildlife management and have important public health and economic implications. © 2020 The Wildlife Society.     

Behavioral responses of male elk to hunting risk

Prey respond to predation risk with a range of behavioral tactics that can vary based on space use and hunting mode of the predator. Unlike other predators, human hunters are often more spatially and temporally restricted, which creates a period of short-duration, high-intensity predation risk for prey. Consequently, identifying the roles different hunting modes (i.e., archery and rifle), hunts for targeted and non-targeted species, and landscape features play in altering spatial and temporal responses of prey to predation risk by humans is important for effective management of harvested populations. From 2009 to 2016, we used a large-scale experiment including 50 animal-years of location data from 38 unique male elk (Cervus canadensis) to quantify changes in movement and resource selection in response to hunters during 3 separate 5-day controlled hunts for antlered males (elk archery, deer [Odocoileus spp.] rifle, and elk rifle) at the Starkey Experimental Forest and Range in northeast Oregon, USA. We evaluated competing hypotheses regarding elk responses to varying levels of prey risk posed by the different hunt types. We predicted that the strength of elk behavioral responses would increase with perceived hunter lethality (i.e., weak response to elk archery but similar response to elk and deer rifle hunts) and that prey response would be closely associated with hunter activity within the diel cycle (greater during diurnal than nocturnal hours) and across hunting seasons. Elk responses were strongest during diurnal hours when hunters were active on the landscape and were generally more pronounced during both rifle hunts than during the archery hunt (supporting our perceived lethality hypothesis). Male elk avoided open roads across all periods except during nocturnal hours of the breeding season and alternated between avoidance of areas with high canopy cover during nocturnal hours and selection during diurnal hours. In combination these patterns led to distinct distributional changes of male elk from pre-hunt to hunt periods. Patterns of male elk selection highlight the importance of managing for heterogeneous landscapes to meet a variety of habitat, harvest, hunter satisfaction, and escapement objectives.     

Habitat Selection by Female Elk During Minnesota's Agricultural Season

Understanding space-use patterns by elk (Cervus canadensis) is essential to alleviating human-elk conflicts, particularly when crop depredation by elk can make it harder to justify to the public the need to restore elk populations to regions with agricultural landscapes. In 2016–2017, we used global positioning system data from 20 female elk to investigate their selection for agricultural cover during the agricultural season (1 May–31 Oct) in northwestern Minnesota, USA. We estimated resource selection functions with resource availability defined at the home range scale. Elk space use was primarily determined by distance to forest cover in areas proximate to agricultural fields. During diurnal periods, elk selected areas with forest cover near agricultural fields planted with legumes and cereal. During nocturnal periods, elk selected for agricultural fields with little to no canopy and that were planted with legumes. We suggest that management of elk in northwestern Minnesota will require practices that discourage the use of agriculture by elk while improving natural habitats within areas managed for elk restoration. We suggest that forestry practices (i.e., thinning and burning) could improve cover and forage openings for elk in restoration areas. Furthermore, managers could work with agricultural producers where elk occur to plant crops favored by elk (i.e., legumes) outside known home ranges and plant fields within home ranges with crops that elk avoided (i.e., hay). Collectively, these practices may shift the ranges of elk herds into restoration areas and lower conflict between the public and elk restoration efforts. © 2020 The Wildlife Society.     

Effects of hunter access and habitat security on elk habitat selection in landscapes with a public and private land matrix

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.     

A Review of Environmental Factors Affecting Elk Winter Diets

Abstract: Decades of research have produced substantial data on elk (Cervus elaphus) diets in winter, when foraging conditions are most likely to affect population dynamics. Using data from 72 studies conducted in western North America between 1938 and 2002, we collated data on elk diets and environmental variables. We used these data to quantify diet selection by elk and to test whether variation in elk diets is associated with habitat type, winter severity, period of winter, human hunting, and study method. Graminoids (grasses and grass-like plants such as sedges) dominated elk diets and consistently occurred at a higher proportion in the diet than in elk foraging habitats, indicating preference. Forbs commonly made up ≤5% of the diet, with no evidence for preference; we conclude that forb use is largely incidental to grazing for graminoids. Browse was consumed in proportion to its availability, implying that the amount of browse in the diet was primarily determined by habitat use rather than selection. Comparing the diets of elk and sympatric ruminants, elk consistently selected graminoids more strongly than sympatric ruminants with the exception of bison (Bison bison), suggesting that elk are not environmentally forced to adopt the graminoid-biased diet that they normally select. The proportion of open meadows and grasslands on winter ranges was strongly and positively associated with graminoid consumption by elk. The proportion of graminoids in the diet was significantly lower in elk experiencing severe winter conditions or predation risk from human hunting. The period of winter (early, middle, and late) had only small effects on elk diets, as did the method by which the diet was determined. Overall, variation in elk diets is well-explained by a consistent tendency to select graminoids if available, modified by winter habitat type, predation risk, and winter severity, which can constrain habitat selection and access to grazing opportunities. To fully understand variation in foraging behavior, biologists should recognize these broad patterns when interpreting resource selection data. Managers should recognize that inconspicuous behavioral responses to environmental stimuli can alter the diet in ways that probably carry nutritional consequences.     

Elk parturition site selection at local and landscape scales

Selection of habitat components by ungulates associated with parturition sites varies among and within species depending upon vulnerability to predators, variation in local topography and climate regimes, and the length of time that the maternal–neonatal unit spends at or near the parturition location. We marked 169 parturition locations of elk (Cervus elaphus nelsoni) in western Wyoming using vaginal implant transmitters and evaluated parturition-specific habitat selection at macro- and microhabitat scales using a resource selection function modeling approach. Elk calved in a variety of habitats, yet demonstrated selection at both spatial scales. We found the strongest support for models that incorporated multiple habitat features and focused on topographical and vegetative cover types that provide physical and thermal cover at the macrohabitat scale and for visual cover models at the microhabitat scale. Models based solely on forage availability or quality were least supported at both scales, which may be indicative of a brief occupation of the parturition location or low heterogeneity in the availability of forage resources on parturition ranges. Results of early elk natural history studies may have represented a bias introduced by variable sightability and accessibility of females with calves and a lack of differentiation between calving and neonatal periods. More clearly defining calving site selection and removing biases toward more open habitats where sightability of neonates is greater may be used by wildlife or land managers to improve or protect calving habitats, which is often a stated objective of management actions. The results of this study suggest that microhabitat is more important to elk and that temporal closures over broad areas versus closures focused on specific macrohabitats may be more effective in protecting calving animals. © 2011 The Wildlife Society.     

Elk Responses to Management Hunting and Hazing

Human-wildlife conflicts are widespread around the world and result in property damage, disease spillover, financial loss, and decreased tolerance of wildlife. Increasing elk (Cervus canadensis) populations and land-use changes in the western United States are challenging resource managers tasked with managing conflict. Lethal and non-lethal management actions are commonly used to remove elk from conflict zones where they are not desired. We used radio-collar location data collected from female elk in 2 study areas in Montana, USA, from 2017–2020 to evaluate population- and individual-level responses to management actions (i.e., hunting, hazing) and environmental factors (i.e., weather, season, time of day). First, we used a generalized linear model with a logit link to evaluate the effects of hunting, hazing, time period, seasonality, and weather on the proportion of collared elk that used a conflict zone. Second, we used an ordinary linear model to assess the influence of hunting, hazing, seasonality, and weather on the duration of time that individual elk remained away from conflict zones. The proportion of elk using conflict zones was reduced by hunting, modestly reduced by hazing and increasing snowpack for 1 study area, increased at night, and decreased by a seasonal trend across months. The time individual elk remained away from conflict zones increased with the number of hazing events that occurred during an event and showed a modest seasonal trend increasing across months. For 1 study area, time away increased with the number of hunting days during an event and increasing snowpack, but the increase was biologically trivial. Our results indicate mixed responses of elk to hunting and hazing actions and provide evidence that management actions can influence elk use of conflict areas. Agencies trying to reduce conflicts may want to consider a combination of hunting and hazing, while accounting for site-specific characteristics to keep elk away from conflict zones. © 2021 The Wildlife Society.     

Mallard resource selection trade‐offs in a heterogeneous environment during autumn and winter

Animals select resources to maximize fitness but associated costs and benefits are spatially and temporally variable. Differences in wetland management influence resource availability for ducks and mortality risk from duck hunting. The local distribution of the Mallard (Anas platyrhynchos) is affected by this resource heterogeneity and variable risk from hunting. Regional conservation strategies primarily focus on how waterfowl distributions are affected by food resources during the nonbreeding season. To test if Mallard resource selection was related to the abundance of resources, risks, or a combination, we studied resource selection of adult female Mallards during autumn and winter. We developed a digital spatial layer for Lake St. Clair, Ontario, Canada, that classified resources important to Mallards and assigned these resources a risk level based on ownership type and presumed disturbance from hunting. We monitored 59 individuals with GPS back‐pack transmitters prior to, during, and after the hunting season and used discrete choice modeling to generate diurnal and nocturnal resource selection estimates. The model that classified available resources and presumed risk best explained Mallard resource selection strategies. Resource selection varied within and among seasons. Ducks selected for federal, state and private managed wetland complexes that provided an intermediate or relatively greater amount of refuge and foraging options than public hunting areas. Across all diel periods and seasons, there was selection for federally managed marshes and private supplemental feeding refuges that prohibited hunting. Mallard resource selection demonstrated trade‐offs related to the management of mortality risk, anthropogenic disturbances, and foraging opportunities. Understanding how waterfowl respond to heterogeneous landscapes of resources and risks can inform regional conservation strategies related to waterfowl distribution during the nonbreeding season.     

Influence of Predator Harvest,Biological Factors,and Landscape on Elk Calf Survival in Idaho

ABSTRACT We evaluated survival of elk (Cervus elaphus) calves on 2 contrasting study areas in north-central Idaho, USA, from 1997 to 2004. Recruitment was modest (>30 calves:100 F [calves of either sex: F elk 1 yr old]) and stable on the South Fork study area and low (<20 calves:100 F) and declining on the Lochsa study area. The primary proximate cause of calf mortality on both study areas was predation by black bears (Ursus americanus) and mountain lions (Puma concolor). We experimentally manipulated populations of black bears and mountain lions on a portion of each study area. Black bear harvest (harvest density/600km²) initially doubled on the Lochsa treatment after manipulating season bag limits. Mountain lion harvest also increased by 60% but varied widely during the manipulation period. Harvest seasons were closed for black bears and mountain lions on the treatment portion of the South Fork study area. Using the Andersen—Gill formulation (A-G) of the Cox proportional hazards model, we examined effects of landscape structure, predator harvest levels, and biological factors on summer calf survival. We used Akaike's Information Criterion (AIC_c) and multimodel inference to assess some potentially useful predictive factors relative to calf survival. We generated risk ratios for both the best models and for model-averaged coefficients. Our models predicted that calf survival was influenced by biological factors, landscape surrounding calf locations, and predator harvest levels. The model that best explained mortality risk to calves on the Lochsa included black bear harvest (harvest density/600 km²), estimated birth mass of calves, and percentage of shrub cover surrounding calf locations. Incorporating a shrub X time interaction allowed us to correct for nonproportionality and detect that effect of shrub cover was only influential during the first 14 days of a calf's life. Model-averaging indicated that estimated birth mass of calves and black bear harvest were twice as important as the next variables, but age of calves at capture was also influential in calf survival. The model that best explained mortality risk to calves on the South Fork included black bear harvest, age of calves at capture, and gender of calves. Model-averaging indicated that age at capture and black bear harvest were twice as important as the next variable, forest with 33–66% canopy cover (Canopy 33–66). Risk to calves decreased when calves occupied areas with more of this forest cover type. Model-averaging also indicated that increased mountain lion harvest lowered calf mortality risk 4% for every 1-unit increase in lion harvest (harvest density/600 km²) but was lower (<25%) in importance compared to age at capture and black bear harvest. Our results suggest that levels of predator harvest, and presumably predator density, resource limitations expressed through calf birth mass, and habitat structure had substantial effects on calf survival. Our results can be generalized to other areas where managers are dealing with low calf elk recruitment. However, because factors vary spatially, a single management strategy applied in different areas will probably not have the same effect on calf survival.     

Targeting hunter distribution based on host resource selection and kill sites to manage disease risk

Endemic and emerging diseases are rarely uniform in their spatial distribution or prevalence among cohorts of wildlife. Spatial models that quantify risk‐driven differences in resource selection and hunter mortality of animals at fine spatial scales can assist disease management by identifying high‐risk areas and individuals. We used resource selection functions (RSFs) and selection ratios (SRs) to quantify sex‐ and age‐specific resource selection patterns of collared (n = 67) and hunter‐killed (n = 796) nonmigratory elk (Cervus canadensis manitobensis) during the hunting season between 2002 and 2012, in southwestern Manitoba, Canada. Distance to protected area was the most important covariate influencing resource selection and hunter‐kill sites of elk (AICw = 1.00). Collared adult males (which are most likely to be infected with bovine tuberculosis (Mycobacterium bovis) and chronic wasting disease) rarely selected for sites outside of parks during the hunting season in contrast to adult females and juvenile males. The RSFs showed selection by adult females and juvenile males to be negatively associated with landscape‐level forest cover, high road density, and water cover, whereas hunter‐kill sites of these cohorts were positively associated with landscape‐level forest cover and increasing distance to streams and negatively associated with high road density. Local‐level forest was positively associated with collared animal locations and hunter‐kill sites; however, selection was stronger for collared juvenile males and hunter‐killed adult females. In instances where disease infects a metapopulation and eradication is infeasible, a principle goal of management is to limit the spread of disease among infected animals. We map high‐risk areas that are regularly used by potentially infectious hosts but currently underrepresented in the distribution of kill sites. We present a novel application of widely available data to target hunter distribution based on host resource selection and kill sites as a promising tool for applying selective hunting to the management of transmissible diseases in a game species.     

Traffic Volume Alters Elk Distribution and Highway Crossings in Arizona

ABSTRACT We used 38,709 fixes collected from December 2003 through June 2006 from 44 elk (Cervus elaphus) fitted with Global Positioning System collars and hourly traffic data recorded along 27 km of highway in central Arizona, USA, to determine how traffic volume affected elk distribution and highway crossings. The probability of elk occurring near the highway decreased with increasing traffic volume, indicating that elk used habitat near the highway primarily when traffic volumes were low (<100 vehicles/hr). We used multiple logistic regression followed by model selection using Akaike's Information Criterion to identify factors influencing probability of elk crossings. We found that increasing traffic rates reduced the overall probability of highway crossing, but this effect depended on both season and the proximity of riparian meadow habitat. Elk crossed highways at higher traffic volumes when accessing high quality foraging areas. Our results indicate that 1) managers assessing habitat quality for elk in areas with high traffic-volume highways should consider that habitat near highways may be utilized at low traffic volumes, 2) in areas where highways potentially act as barriers to elk movement, increasing traffic volume decreases the probability of highway crossings, but the magnitude of this effect depends on both season and proximity of important resources, and 3) because some highway crossings still occurred at the high traffic volumes we recorded, increasing traffic alone will not prevent elk-vehicle collisions. Managers concerned with elk-vehicle collisions could increase the effectiveness of wildlife crossing structures by placing them near important resources, such as riparian meadow habitat.     

The Restoration of Elk (Cervus elaphus) in Ontario, Canada: 1998–2005

In 1997, a plan to restore Elk (Cervus elaphus) to Ontario was approved by the provincial government. The objective of the Ontario elk restoration program, a multipartnered collaboration, was to restore a species that had been extirpated from the province during the 1800s. During 1998–2001, 460 elk were acquired from Elk Island National Park, Alberta, for release in four areas of Ontario. As greater than 90% of the elk were radio collared, monitoring provided detailed information on the dynamics of the four populations. Comprehensive research projects using graduate students were implemented to determine the environmental impact of releasing elk in Ontario. Those studies are in progress or have been completed and include the effect of wolf predation on restored elk, white‐tailed deer and elk resource overlap, the development of genetic profiles for elk, and solutions for elk/human conflicts. Mortality of the released elk averaged 41% (190/460) during 1998–2004 with annual mortality generally declining over time in each release area. The primary causes of elk mortality included wolf predation (25% of mortalities), illegal shooting (13%), stress‐related emaciation (13%) (partially due to the stress of relocation), bacterial infections (7%), and collisions with vehicles (6%). Productivity has been high in one of the release areas with 24–65% of the cows being observed with calves during late winter surveys. However, productivity has been low in two of the northern release areas due to a variety of factors including wolf predation. In some areas, dispersion of elk appeared to be related to the length of time animals were kept in pens prior to release. The precalving population estimate for Ontario in March 2004 was 375–440 elk. A comprehensive program review was conducted in 2003/2004 that included recommendations relating to the future management of elk in Ontario.     

Hiding Without Cover? Defining Elk Security in a Beetle-Killed Forest

Recent mountain pine beetle (Dendroctonus ponderosae; pine beetle) outbreaks in the western United States have affected nearly 18 million ha of pine (Pinus spp.) forest and are unprecedented in spatial extent, severity, and duration, yet little is known about wildlife responses to large-scale insect outbreaks. Elk (Cervus canadensis) are important wildlife whose dominant management paradigm on public lands has focused on providing security habitat to increase survival during hunting seasons and to maintain elk presence on public lands to promote hunter opportunity. To assess the effect of pine beetles and associated changes in forest structure on elk security, we used a time series to characterize canopy cover pre- and post-pine beetle outbreak, characterized relative canopy cover among the dominant forest types in the study area post-pine beetle outbreak, and used global positioning system location data from male and female elk to define habitat relationships and security during the archery and rifle hunting seasons. Our study area was within the Elkhorn Mountains of southwest Montana, USA, 2015–2017, which experienced 80% mortality of lodgepole pine (Pinus contorta) forests during a pine beetle outbreak that peaked in 2008. We observed an 8.5% reduction in canopy cover within pine beetle-infested lodgepole pine forests, yet canopy cover remained relatively high among other forest types post-outbreak. The top-ranked habitat security models contained positive relationships with canopy cover, distance to motorized routes, terrain ruggedness, and slope with few notable differences among sexes and seasons. Across sexes and seasons, 75% and 50% of elk use was within areas with average canopy cover values ≥31 ± 6.65 (SD)% and ≥53 ± 5.7% that were an average of ≥2,072 ± 187.93 m and ≥3,496 ± 157.32 m from a motorized route, respectively. Therefore, we recommend fall elk security be defined as areas that meet these criteria for minimum canopy cover and distance from motorized routes in the Elkhorn Mountains and in other landscapes with similar forest characteristics and hunting pressures. Although we observed expected reductions in canopy cover within pine beetle-infested forests, defoliation alone did not appear to negatively affect elk security or reduce canopy cover below our management recommendations. Nonetheless, because of the prevalence of standing dead trees in our study area, we recommend future work that investigates the relationships with pine beetle-infested areas post-blowdown because changes in ground structure and costs of locomotion may affect elk habitat and security. © 2019 The Authors. Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Mothers' Movements: Shifts in Calving Area Selection by Partially Migratory Elk

Loss of migratory behavior or shifts in migratory ranges are growing concerns to wildlife managers. How ungulates prioritize safety from predators at the expense of high-quality foraging opportunities during calving may be key to understanding these shifts and long-term reproductive success. We compared trade-offs in selection for forage and predation risk by elk (Cervus canadensis) following 3 migratory tactics (western and eastern migration and resident) during 2 time periods in a declining (by almost 70% from 2002–2016), partially migratory elk population adjacent to Banff National Park in Alberta, Canada. We hypothesized that regardless of migratory tactic, maternal elk would show stronger trade-offs between high-quality foraging associated with higher predation risk and forage resources of lower-quality yet lower risk on calving ranges than on ranges used during summer because of vulnerability of their offspring. Additionally, we hypothesized these trade-offs would occur at high (2002–2006) and low (2013–2016) elk population sizes. We used a machine-learning algorithm to predict dates of parturition based on global positioning system (GPS) movements of elk equipped with vaginal implants (n = 60) and predictions were within 1.43 ± 0.85 (SE) days of the known date. We applied the model to an additional 58 GPS-collared elk without vaginal implants. Based on changes in localized movements, we defined calving areas as the 26 days post-parturition and compared habitat characteristics of calving areas to 10 similar-sized areas centered on random locations during summer for the same individual in a latent selection framework. Across the 2 time periods, parturition occurred from 8 May–11 July with median parturition dates differing among migratory tactics and residents shifting towards an earlier parturition date in the later period. All elk, regardless of migratory tactic and time period, selected calving areas with greater forage resources than were available on areas used during summer, with no evidence for greater selection of areas that reduced predation risk at the expense of higher-quality foraging. Calving season selection for areas with abundant forage exposed western migrants to high risk of bear (Ursus spp.) predation, residents to high risk of wolf (Canis lupus) predation, and eastern migrants to low risk of bear and wolf predation. Patterns in exposure to predation risk during calving between migratory tactics were consistent with the recent decline in western migrants and increase in eastern migrants, implying that conditions on calving areas contributed to observed changes in the number of elk following these tactics. © 2021 The Wildlife Society.     

Identifying Sites for Elk Restoration in Arkansas

Abstract: We used spatial data to identify potential areas for elk (Cervus elaphus) restoration in Arkansas. To assess habitat, we used locations of 239 elk groups collected from helicopter surveys in the Buffalo National River area of northwestern Arkansas, USA, from 1992 to 2002. We calculated the Mahalanobis distance (D²) statistic based on the relationship between those elk-group locations and a suite of 9 landscape variables to evaluate winter habitat in Arkansas. We tested model performance in the Buffalo National River area by comparing the D² values of pixels representing areas with and without elk pellets along 19 fixed-width transects surveyed in March 2002. Pixels with elk scat had lower D² values than pixels in which we found no pellets (logistic regression: Wald χ² = 24.37, P < 0.001), indicating that habitat characteristics were similar to those selected by the aerially surveyed elk. Our D² model indicated that the best elk habitat primarily occurred in northern and western Arkansas and was associated with areas of high landscape heterogeneity, heavy forest cover, gently sloping ridge tops and valleys, low human population density, and low road densities. To assess the potential for elk-human conflicts in Arkansas, we used the analytical hierarchy process to rank the importance of 8 criteria based on expert opinion from biologists involved in elk management. The biologists ranked availability of forage on public lands as having the strongest influence on the potential for elk-human conflict (33%), followed by human population growth rate (22%) and the amount of private land in row crops (18%). We then applied those rankings in a weighted linear summation to map the relative potential for elk-human conflict. Finally, we used white-tailed deer (Odocoileus virginianus) densities to identify areas where success of elk restoration may be hampered due to meningeal worm (Parelaphostrongylus tenuis) transmission. By combining results of the 3 spatial data layers (i.e., habitat model, elk-human conflict model, deer density), our model indicated that restoration sites located in west-central and north-central Arkansas were most favorable for reintroduction.     

How Size and Condition Influence Survival and Cause-Specific Mortality of Female Elk

The size of animal populations fluctuates with number of births, rate of immigration, rate of emigration, and number of deaths. For many ungulate populations, adult female survival is the most important factor influencing population growth. Therefore, increased understanding of survival and causes of mortality for adult females is fundamental for conservation and management. The objectives of our study were to quantify survival rates of female elk (Cervus canadensis) and determine cause-specific mortality. We predicted that hunter harvest would be the leading cause of mortality. Further, we predicted that hunters would harvest animals that were in prime age (2–9 yr) and in better condition than elk predated by mountain lions (Puma concolor). From 2015 to 2017, we captured 376 female elk in central Utah, USA. We assessed body size and condition of captured elk, fitted each animal with a global positioning system-collar, and determined cause of death when we received mortality signals. We estimated survival using Kaplan-Meier estimates and Cox proportional hazard models within an Akaike's Information Criterion model selection framework to identify covariates that influenced survival. We analyzed differences in size and condition measurements between harvested elk and predated elk using analysis of variance tests. Our best model indicated consistent survival across years; mean survival was 78.3 ± 3.5% (SE) including hunter harvest and 95.5 ± 1.7% without hunter harvest. In decreasing order of importance, elk mortality occurred from hunter harvest (21.2%), mountain lion predation (3.7%), depredation removal (0.5%), automobile collision (0.3%), disease (0.3%), complications during calving (0.3%), and those characterized as undetermined (1.3%). Neck circumference and body length were negatively associated with survival, suggesting that larger animals in good condition had lower survival as a result of hunter harvest. Individuals that died because of cougar predation were smaller and had less loin muscle than the average animal. Hunters removed large, healthy, prime-aged females, individuals that likely have a greater effect on population growth than elk lost to other predators. If the proportion of larger, healthy females in the population begins to decline, hunting practices may require adjustment because hunters may be removing individuals with the greatest reproductive value. © 2021 The Wildlife Society.     

Behavioral Responses of North American Elk to Recreational Activity

ABSTRACT Off-road recreation on public lands in North America has increased dramatically in recent years. Wild ungulates are sensitive to human activities, but the effect of off-road recreation, both motorized and nonmotorized, is poorly understood. We measured responses of elk (Cervus elaphus) to recreational disturbance in northeast Oregon, USA, from April to October, 2003 and 2004. We subjected elk to 4 types of recreational disturbance: all-terrain vehicle (ATV) riding, mountain biking, hiking, and horseback riding. Motion sensors inside radiocollars worn by 13 female elk recorded resting, feeding, and travel activities at 5-minute intervals throughout disturbance and control periods. Elk fed and rested during control periods, with little time spent traveling. Travel time increased in response to all 4 disturbances and was highest in mornings. Elk travel time was highest during ATV exposure, followed by exposure to mountain biking, hiking, and horseback riding. Feeding time decreased during ATV exposure and resting decreased when we subjected elk to mountain biking and hiking disturbance in 2003. Our results demonstrated that activities of elk can be substantially affected by off-road recreation. Mitigating these effects may be appropriate where elk are a management priority. Balancing management of species like elk with off-road recreation will become increasingly important as off-road recreational uses continue to increase on public lands in North America.     

Contrasting Effects of Wolves and Human Hunters on Elk Behavioral Responses to Predation Risk

ABSTRACT 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.