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Large herbivore populations respond strongly to remotely sensed measures of primary productivity. Whereas most studies in seasonal environments have focused on the effects of spring plant phenology on juvenile survival, recent studies demonstrated that autumn nutrition also plays a crucial role. We tested for both direct and indirect (through body mass) effects of spring and autumn phenology on winter survival of 2315 mule deer fawns across a wide range of environmental conditions in Idaho, USA. We first performed a functional analysis that identified spring and autumn as the key periods for structuring the among-population and among-year variation of primary production (approximated from 1 km Advanced Very High Resolution Radiometer Normalized Difference Vegetation Index (NDVI)) along the growing season. A path analysis showed that early winter precipitation and direct and indirect effects of spring and autumn NDVI functional components accounted for 45% of observed variation in overwinter survival. The effect size of autumn phenology on body mass was about twice that of spring phenology, while direct effects of phenology on survival were similar between spring and autumn. We demonstrate that the effects of plant phenology vary across ecosystems, and that in semi-arid systems, autumn may be more important than spring for overwinter survival.

For many species, behavioral modification is an effective strategy to mitigate negative effects of harsh and unpredictable environmental conditions. When behavioral modifications are not sufficient to mitigate extreme environmental conditions, intrinsic factors may be the primary determinant of survival. We investigated how movement behavior, and internal (i.e., nutrition and age) and external (i.e., food availability and snow depth) states affect survival over winter of a long‐lived and highly faithful species (mule deer; Odocoileus hemionus). We first tested whether animals changed their behavior during winter based on internal and external states; we subsequently investigated how behavior and state interacted to influence survival in the face of extraordinary winter conditions. Movement behavior changed minimally as a function of age and nutrition; yet, movement behavior affected survival—animals that exhibited more restricted movements were more likely to succumb to mortality overwinter than animals with less restricted movements. Additionally, nutrition and cumulative snow depth had a strong effect on survival: animals that were exposed to deep snow and began winter with low fat were much less likely to survive. Behavior was an effective tool in securing survival during mild or moderate winters, but nutrition ultimately underpinned survival during harsh winters.

Quantifying the relative influence of multiple mechanisms driving recent range expansion of non‐native species is essential for predicting future changes and for informing adaptation and management plans to protect native species. White‐tailed deer (Odocoileus virginianus) have been expanding their range into the North American boreal forest over the last half of the 20th century. This has already altered predator–prey dynamics in Alberta, Canada, where the distribution likely reaches the northern extent of its continuous range. Although current white‐tailed deer distribution is explained by both climate and human land use, the influence each factor had on the observed range expansion would depend on the spatial and temporal pattern of these changes. Our objective was to quantify the relative importance of land use and climate change as drivers of white‐tailed deer range expansion and to predict decadal changes in white‐tailed deer distribution in northern Alberta for the first half of the 21st century. An existing species distribution model was used to predict past decadal distributions of white‐tailed deer which were validated using independent data. The effects of climate and land use change were isolated by comparing predictions under theoretical “no‐change between decades” scenarios, for each factor, to predictions under observed climate and land use change. Climate changes led to more than 88%, by area, of the increases in probability of white‐tailed deer presence across all decades. The distribution is predicted to extend 100 km further north across the northeastern Alberta boreal forest as climate continues to change over the first half of the 21st century. Quantifying the relative influence of multiple mechanisms driving recent range expansion of non‐native species is essential for predicting future changes and for informing adaptation and management plans to protect native species. Our objective was to quantify the relative importance of land use and climate change as drivers of white‐tailed deer range expansion and to predict decadal changes in northern white‐tailed deer distribution for the first half of the 21st century. Climate changes led to more than 88%, by area, of the increases in probability of white‐tailed deer presence across all decades. The distribution is predicted to extend 100 km further north across the northeastern Alberta boreal forest as climate continues to change over the first half of the 21st century.

In many southern boreal ecosystems of North America, wolves are the primary predators of white‐tailed deer, and white‐tailed deer are the primary prey of wolves. Furthermore, wolf–deer systems have and will continue to become more common as white‐tailed deer range continues expanding northward in North America. Despite this, there is little information on kill rates of wolves on deer (i.e., the number of deer killed per wolf per unit of time)—a fundamental metric of wolf predation on deer—and how kill rates vary with deer density, wolf density, and environmental conditions. We estimated kill rates of wolves on deer before, during, and after a historically mild winter in the Greater Voyageurs Ecosystem, Minnesota, USA. Kill rates of wolves on deer were low (0.009–0.018 deer/wolf/day) in fall, peaked in February (0.050 deer/wolf/day), and quickly declined to 0 deer/wolf/day by April. The kill rates of wolves on deer we observed in winter were some of the lowest kill rates of wolves on deer that have been documented. Wolves in the Greater Voyageurs Ecosystem appeared unable to catch and kill a sufficient number of deer to meet their daily energetic requirements during Winter 2023–2024, and thus most wolves likely lost weight during winter, a period when wolves are typically in peak physical condition. The rates of wolf predation we observed appeared to be well below those needed to decrease deer population density in the GVE. Thus, our work, in combination with numerous other studies, indicates winter conditions are the primary driver of deer population change in northern climates. We estimated kill rates of wolves on deer before, during, and after a historically mild winter in the Greater Voyageurs Ecosystem, Minnesota, USA. Kill rates of wolves on deer were low in fall, peaked in February, and quickly declined to 0 deer/wolf/day by April. Wolves in the Greater Voyageurs Ecosystem appeared unable to catch and kill a sufficient number of deer to meet their daily energetic requirements during Winter 2023–2024, and thus most wolves likely lost weight during winter, a period when wolves are typically in peak physical condition.

A relationship between winter weather and survival of northern ungulates has long been established, yet the possible roles of biological (e.g., nutritional status) and environmental (e.g., weather) conditions make it important to determine which potential limiting factors are most influential. Our objective was to examine the potential effects of individual (body mass and age) and extrinsic (winter severity and snowmelt conditions) factors on the magnitude and timing of mortality for adult (>2.5 years old) female white‐tailed deer (Odocoileus virginianus [Zimmerman, 1780]) during February–May in the Upper Peninsula of Michigan, USA. One hundred and fifty deer were captured and monitored during 2009–2015 in two areas with varying snowfall. February–May survival ranged from 0.24 to 0.89 (mean = 0.69) across years. Mortality risk increased 1.9% with each unit increase in cumulative winter severity index, decreased 8.2% with each cumulative snow‐free day, and decreased 4.3% with each kg increase in body mass. Age and weekly snow depth did not influence weekly deer survival. Predation, primarily from coyote (Canis latrans [Say, 1823]) and wolves (Canis lupus [L., 1758]), accounted for 78% of known‐cause mortalities. Our results suggest that cumulative winter severity, and possibly to a lesser degree deer condition entering winter, impacted deer winter survival. However, the timing of spring snowmelt appeared to be the most influential factor determining late‐winter mortality of deer in our study. This supports the hypothesis that nutrition and energetic demands from weather conditions are both important to northern ungulate winter ecology. Under this model, a delay of several weeks in the timing of spring snowmelt could exert a large influence on deer survival, resulting in a survival bottleneck. Northern ungulate population dynamics are correlated with winter weather patterns, and our goal was to evaluate when and why white‐tailed deer winter mortality occurs. We modeled weekly mortality risk of 150 adult female white‐tailed deer in Michigan in response to environmental and biological factors. The results suggested that body mass, severity of winter weather, and timing of spring snowmelt are influential on deer mortality, with the timing of snowmelt explaining the greatest amount of variation.

The importance of conserving migratory populations is recognized across a variety of ungulate taxa, yet the demographic benefits of migration remain uncertain for ungulate populations that exhibit partial migration. We hypothesized that migratory pronghorn (Antilocapra americana) would experience greater survival compared to residents by moving longer distances to avoid severe winter weather and access higher quality forage. We used a Bayesian time-to-event approach to analyze the fates of 175 radio-collared adult female pronghorn monitored over 8 biological years (2004–2011) in the Northern Sagebrush Steppe ecosystem. Annual survivorship of migratory pronghorn was 7% higher on average compared to residents but not statistically different. Migratory pronghorn had higher survivorship in summer and winter compared to residents, and few mortalities were observed during the short autumn and spring migration periods. Mortality risk for both movement tactics intensified under more severe winter weather; winter weather severity alone best explained annual pronghorn mortality risk. The top model predicted survival rates to decline on average by 56% over the range of observed winter climatic conditions. To minimize human impacts to pronghorn during extreme climatic events, we recommend working with transportation departments and land managers to enhance pronghorn crossings of roads and railroads, and landholders to modify fences to wildlife-friendly standards.

Elk (Cervus canadensis) are high-profile game animals for many states in the western United States, yet over the past several decades some populations have experienced a persistent and broad-scale decline in recruitment. Over this same period, gray wolves (Canis lupus) have become an integral component of many western landscapes and agencies are increasingly challenged to maximize hunting opportunities of ungulates via predator management while simultaneously ensuring wolf conservation. To better understand the implications of predator management on elk populations, we monitored survival of 1,244 adult female elk and 806 6-month-old calves from 29 populations distributed throughout Idaho, USA, from 2004 to 2016. We developed predictive models of mortality that related mortality risk to wolf pack size, winter conditions, and individual-level characteristics. Annual mortality rates (excluding harvest) for adult females and calves were 0.09 and 0.40, respectively. Calf mortality was predicted best with a model that included additive effects of chest girth at time of capture, mean size of surrounding wolf packs, and snow depth. Adult female mortality was predicted best with a model that included female age, mean size of surrounding wolf packs, and snow depth. Based on a sensitivity analysis, chest girth had the largest effect on risk of mortality for calves followed by pack size and snow depth. Other than the effect of senescence in the oldest (>15 yr) individuals, pack size and snow depth had the largest effect on risk of mortality for adult females. We estimated cause-specific mortality and predation was the dominant cause of known-fate mortalities for adult females (35% mountain lion [Puma concolor] and 32% wolf) and calves (45% mountain lion and 28% wolf), whereas malnutrition accounted for 9% and 10% of adult female and calf mortalities, respectively. Wolves preferentially selected smaller calves and older adult females, whereas mountain lions showed little preference for calf size or age class of adult females. Our study indicates managers can increase elk survival by reducing wolf pack sizes on surrounding winter ranges, especially in areas where, or during years when, snow is deep. Additionally, managers interested in improving over-winter calf survival can implement actions to increase the size of calves entering winter by increasing the nutritional quality of summer and early fall forage resources. Although our study was prompted by management questions related to wolves, mountain lions killed more elk than wolves and differences in selection of individual elk indicate mountain lions may have comparably more of an effect on elk population dynamics. Although we were unable to relate changes in mountain lion populations to elk survival in our study, future research should seek a better understanding of multi-predator systems, including how management of one predator affect others and ultimately how these interactions affect elk survival.

Climate change is an increasingly important driver of biodiversity loss. The ectothermic nature of amphibians may make them particularly sensitive to changes in temperature and precipitation regimes, adding to declines from other threats. While active season environmental conditions can influence growth and survival, effects of variation in winter conditions on population dynamics are less well-studied. Given that extreme winter temperatures can influence amphibian survival and fitness, we expected that increased winter severity—as measured by variability in winter temperatures and snow cover—would be associated with decreased occupancy, and that populations that experience more severe winters would have the largest sensitivities and show the greatest declines.

Variation among demographic rates for a population reflects the allocation of available energy by individuals to competing life‐history strategies. Species exhibiting slow‐paced life histories often prioritize energy allocation to adult survival over any single reproductive event, therefore maximizing future reproductive potential. Survival of adult female ungulates is generally high with little variability, whereas survival of young is lower and often highly variable. When adult survival is high with low variability, juvenile survival may have a proportionally greater effect on population growth or decline. Weather also may affect population dynamics directly by influencing survival of young or adults, or indirectly through changes in nutritional condition of adult females that influence population growth rates. We experimentally manipulated forage availability during winter, by supplementing native forage with high‐energy pelleted feed ad libitum, to a subset of a population of mule deer (Odocoileus hemionus) to understand the effects of winter nutrition on survival of adult females and their young born the subsequent summer. We evaluated the effects of winter nutrition, individual‐based parameters, and environmental covariates on survival of adult female mule deer from 2013 to 2018, and neonatal mule deer from 2014 to 2016. We documented a 26% decrease in annual survival of adult female mule deer in 2017 in response to increased snowpack during the preceding winter. Neonates born to females that receive enhanced nutrition during winter preceding parturition had higher survival to weaning (0.49, SE = 0.12), compared to neonates born to females that did not receive enhanced nutrition (0.29, SE = 0.07). We observed no effect of enhanced winter nutrition on survival of adult females. Our results suggested winter nutrition of maternal females may influence juvenile survival and demonstrates the importance of forage quality available to adult females during mid‐pregnancy. Although we were unable to detect an effect of winter forage on survival of adults, direct effects of deep winter snow resulted in lower survival of adult females. Low survival of adult females in our study population is indicative of a declining population.

The North American black bear (Ursus americanus) is an opportunistic omnivore that depends on seasonal availability of fruits, nuts, grasses, and forbs for survival. Black bears on the urban–wildland interface also use anthropogenic food resources, especially when natural food resources are scarce. Consequently, natural food failure can exacerbate human–bear conflict, resulting in increases in human‐caused mortality via vehicle strikes and management removal. Climate change is expected to increase the frequency and severity of extreme weather events, including drought or late frost. These climatic events may affect the spring growth resulting in loss of natural foods for bears and lead to heightened human–bear conflict in the future. In this study, we examined the effects of weather (snowpack and final freeze dates) on natural survival and cause‐specific mortality (management removal and vehicle collision) of black bears in northwestern Nevada, using an extensive capture–recapture database (509 bears captured between 1998 and 2022). Our results indicated that late freeze dates were associated with a higher probability of conflict, increased probability of management removal, and reduced natural survival. Snowpack (snow‐water equivalent) was weakly correlated with the probability of management removal, but the association was much weaker than the effect of late freeze dates. Anticipating the effects of late frost and snowpack on human–black bear conflict will help managers better anticipate and respond to potential high‐conflict events.