Timing of grizzly bear (Ursus arctos) parturition during hibernation has been explained by ancestral traits (delayed implantation, altricial young, obligate maternal denning), but the ultimate driver underlying precise timing has not been fully explored. Capitalizing on an observed latitudinal increase in denning duration among four populations in interior North America, we tested two alternative hypotheses. First, that birth timing results from a physiological cue that synchronizes implantation with the onset of hibernation, allowing females to forgo reproduction should they lack adequate fat stores. Alternatively, that parturition is optimally timed relative to den exit to balance an energetic tradeoff between minimizing lactation time to protect the mother and maximizing developmental time to increase cub survival. Using parturition dates previously predicted from accelerometer data (27 Dec-28 Feb), we classified 115 females according to apparent litter survival when first visually observed after den exit: 57% successful (with cubs), 22% unsuccessful (alone), and 21% unknown (not observed). The number of days between birth and den exit showed no association with latitude (p = 0.29). It averaged 103 days among successful females but only 77 days among unsuccessful females (p < 0.001) owing to later births and earlier exit. With each increasing degree of latitude, birth date increased by 1.0 and number of days between den entry and birth increased by 2.5 (p < 0.001). Implantation dates were not centered on den entry dates (p < 0.001). These results supported the energetic tradeoff hypothesis and suggested natural selection has favored a consistent number of days between parturition and den exit under average body conditions and shifts toward later or earlier births for females with lower or higher levels of bodily stored energy, respectively. This flexible tradeoff may support resilience to climate change and present a possible mechanism explaining reduced natality and cub survival in high-density populations.

Given the importance of protected areas for biodiversity, the growth of visitation to many areas has raised concerns about the effects of humans on wildlife. In 2020, the COVID-19 pandemic led to temporary closure of national parks in the United States, offering a pseudonatural experiment to tease apart the effects of permanent infrastructure and transient human presence on animals. We compiled GPS tracking data from 229 individuals of 10 mammal species in 14 parks and used third-order hierarchical resource selection functions to evaluate the influence of the human footprint on animal space use in 2019 and 2020. Averaged across all parks and species, animals avoided the human footprint, whether the park was open or closed. However, although animals in remote areas showed consistent avoidance, on average those in more developed areas switched from avoidance to selection when protected areas were closed. Findings varied across species: some responded consistently negatively to the footprint (wolves, mountain goats), some positively (mule deer, red fox) and others had a strong exposure-mediated response (elk, mountain lion). Furthermore, some species responded more strongly to the park closure (black bear, moose). This study advances our understanding of complex interactions between recreation and wildlife in protected areas.

Diel activity rhythms, representing the behavioral pattern of the sleep-wake cycle, may be adjusted by wildlife in response to changes in environmental conditions. An increase in nocturnality is typically recognized as an adaptive strategy to segregate from humans and mitigate heat stress. Numerous studies have investigated spatial patterns and habitat use of large carnivores in human-modified landscapes, but little research has examined their activity rhythms. We compiled Global Positioning System data (2004-2022) for 139 brown bears Ursus arctos from six populations across Europe, representing a human-modified landscape, and the Greater Yellowstone Ecosystem, U.S.A., representing a landscape with limited human impact, which we used to calculate hourly movement rates as an activity proxy. Using a Bayesian approach to model the temporal autocorrelation of activity data, we tested if the extent of nocturnality in brown bears is modulated by intensity of human encroachment, accounting for primary productivity and maximum ambient temperature. All bear populations exhibited a predominantly bimodal, crepuscular pattern of activity, although Yellowstone bears were proportionally more crepuscular and diurnal. Whereas the effect of primary productivity was variable, all European populations became more nocturnal in response to higher human encroachment and reduced diurnal and crepuscular activity at higher summer temperatures, decreasing overall diel activity levels. Yellowstone bears displayed the greatest shift towards nocturnality among all populations in response to increasing human encroachment, and increased nocturnal activity to compensate for lower diurnal and crepuscular activity at higher summer temperatures. Our research indicates that European bears in human-modified landscapes may be reaching a limit in the behavioral plasticity they can manifest in their activity patterns, being already constrained into increased nocturnality. Our findings enhance the understanding of brown bear adaptive capacity to accommodate future changes, such as urbanization and increasing temperatures, to the ecosystems they inhabit.

Plasticity of diel activity rhythms may be a key element for adaptations of wildlife populations to changing environmental conditions. In the last decades, grizzly bears Ursus arctos in the Greater Yellowstone Ecosystem (GYE) have experienced notable environmental fluctuations, including changes in availability of food sources and severe droughts. Although substantial research has been conducted on grizzly bear diets, space use, and demographic parameters, studies on factors that may influence their diel activity patterns are lacking. We investigated diel activity of grizzly bears in the GYE as a function of anthropogenic landscape modification, maximum daily ambient temperature, drought severity, and bear density. Specifically, we used accelerometry readings of 169 bears (39 females, 130 males) from 2009 to 2022 to compute three complementary activity measures, hourly intensity of activity, daily active minutes, and active bout length, each used as a response variable within a Bayesian modeling framework. Grizzly bears generally exhibited bimodal diel activity, with crepuscular peaks and slight variations across seasons. Females with young (i.e. cubs or yearlings) were an exception, with more pronounced diurnal activity patterns, possibly as a strategy to avoid infanticide by dominant males. Landscape modification and maximum ambient temperature were the factors most strongly associated with activity patterns of grizzly bears, with greater nocturnality observed in lone females and males as these factors increased. Females with young were comparatively less affected. The GYE is changing because of increasing land development, human recreation pressures, and effects of climate change. Given their greater diurnal activity compared with other cohorts, female grizzly bears with dependent offspring may be more constrained in their ability to modify activity patterns. Our findings add to a growing body of research emphasizing the importance of the temporal dimension of wildlife behavior as a critical factor in assessing species adaptability and vulnerability in a changing world.

BACKGROUND:Information on reproductive success is crucial to understanding population dynamics but can be difficult to obtain, particularly for species that birth while denning. For grizzly (Ursus arctos) and polar bears (U. maritimus), den visits are impractical because of safety and logistical considerations. Reproduction is typically documented through direct observation, which can be difficult, costly, and often occurs long after den departure. Reproduction could be documented remotely, however, from post-denning movement data if discernable differences exist between females with and without cubs. METHODS:We trained support vector machines (SVMs) with eight variables derived from telemetry data of female grizzly (2000-2022) and polar bears (1985-2016) with or without cubs during seven periods with lengths ranging from 5 to 60 days starting at den departure. We assessed SVM classification accuracy by withholding two samples (one cub-present, one cub-absent), training SVMs with the remaining data, predicting classification of the withheld samples, and repeating this process for each sample combination. Additionally, we evaluated how classification accuracy for grizzly bears was influenced by sample size, length of the post-departure period, and frequency of standardized location estimates. RESULTS:Accuracy of predicting cub presence or absence was 87% for grizzly bears with only 5 days of post-departure data and increased to a maximum of 92% with 20 days of data. For polar bears, accuracy was 86% at 5 days post-departure and increased to a maximum of 93% at 50 days. Classification accuracy for grizzly bears increased from 76 to 90% when sample size increased from 10 to 30 bears while holding period length constant (30 days) but did not increase at larger sample sizes. When sample size was held constant, increasing the length of the post-departure period did not affect classification accuracy markedly. CONCLUSION:Presence or absence of grizzly and polar bear cubs can be identified with high accuracy even when SVM models are trained with limited data. Detecting cub presence or absence remotely could improve estimates of reproductive success and litter survival, enhancing our understanding of factors affecting cub recruitment.

Strategies animals use to navigate human-dominated landscapes frequently mimic anti-predator responses employed by prey species. Understanding how large carnivores respond to outdoor recreation is important for conservation, particularly in protected areas with preservation mandates. Visitation to Yellowstone National Park doubled from 1980 to 2015, increasing the need to examine potential changes in behavior of grizzly bears (Ursus arctos) in relation to human recreation sites (trails, backcountry campsites). We developed integrated step-selection functions to explore how recreation sites influenced the movement rate and selection by male and female grizzly bears. Further, we tested whether time of day (diurnal, crepuscular, nocturnal) and restrictions to human access (i.e., restricted, unrestricted) modified bear responses and then compared behaviors based on proximity to recreation sites. Male grizzly bears used trails to travel during crepuscular and nocturnal hours and exhibited more pronounced behavior in restricted areas compared with unrestricted areas, suggesting recreation in unrestricted areas influenced the behavior of male bears. In contrast, female bears varied their movement rate and selection of trails in restricted areas much more than in unrestricted areas, suggesting females may make security tradeoffs between male bears and people. Both sexes used trails, likely as energetically efficient travel corridors; however, our analyses did not indicate that bears spent time near backcountry campsites. The sex-based differences in selection and movement patterns associated with trails and campsites suggest a single management approach for recreation may not equally benefit all bears. Recreation impacts on wildlife are complex to characterize and predict, but simultaneously modeling movement and selection provides a more comprehensive assessment of strategies animals use to navigate perceived risk.

Grizzly bear (Ursus arctos) feeding on migratory army cutworm moths (Euxoa auxiliaris) was first documented by the Interagency Grizzly Bear Study Team (IGBST) during the early 1980s in the southeastern portion of the Greater Yellowstone Ecosystem (GYE). Since those initial observations, use of this seasonally available food resource by grizzly bears has grown substantially. As of 2023, we have documented 4,754 observations of grizzly bears feeding or digging at high-elevation talus slopes. We used those records to identify 36 unique moth sites in the GYE, assessed geographic characteristics of these sites, and documented chronology and frequency of use by grizzly bears. We used occurrences of radio-collared grizzly bears to identify a sample of bears that foraged at moth sites and investigated characteristics of fidelity, duration, diel activity, and movement patterns. Grizzly bears exhibited high fidelity to specific moth sites within and across years. Bears showed approximately 50% reductions in movement metrics while using sites, albeit with increased activity patterns. Estimates of feeding metrics by females exceeded those of males in intensity and duration of both use and daily activity. Given increasing human impacts and interest in observing this feeding interaction, quantifying metrics that describe the spatial and temporal patterns of moth site use by grizzly bears could be beneficial for future management. Further analyses are needed to fully examine the relationship between caloric influences of moth use on grizzly bear demographic rates.

Expanding human pressure has reduced natural habitats globally and motivated strategies to conserve remaining natural habitats. Decisions about conservation on private lands, however, are typically made by local stakeholders who are motivated by the elements of nature they most highly value. Thus, national prioritization for conservation should be complemented by local analysis of species or habitats that most influence local landowner decisions. We demonstrate within the Greater Yellowstone Ecosystem how quantitative mapping of wildlife species that are highly valued by local residents can be integrated with indices of ecosystem integrity to prioritize private lands for conservation. We found that natural vegetation cover (NVC) comprised 81% of the private lands. Some watersheds have lost 6% of NVC since 2001 and developed lands now cover.40% of their areas. Locations high in ecological value, elk habitat, and grizzly habitat occurred in different biophysical settings. Consequently, only 2% of the NVC supports high levels of all three biodiversity measures and 26% of this area was within conservation easements. The remaining areas of high biodiversity value that are unprotected are priorities for conservation. We suggest that national-scale conservation planning will be most effective on private lands if additional within-ecoregion analyses are done on the elements of biodiversity that are most valued by local people.

Long-term wildlife research and monitoring programs strive to maintain consistent data collections and analytical methods. Incorporating new techniques is important but can render data sets incongruent and limit their potential to discern trends in demographic parameters. Integrated population models (IPMs) can address these limitations by combining data sources that may span different periods into a unified statistical framework while providing a holistic view of population dynamics. We developed an IPM in a Bayesian framework for grizzly bears (Ursus arctos) in the Greater Yellowstone Ecosystem. We coupled demographic data with multiple, independent population count data to link annual changes in abundance with vital rates over 4 decades (1983-2023). Abundance increased threefold from an estimated 270 individuals in 1984 to 1030 individuals in 2023. Parameter estimates indicated survival of bears >= 2 years of age was high, contributing to robust population growth during the 1980s (lambda = 1.023 [50 % interquartile range = 0.993-1.082]) and 1990s (lambda = 1.064 [1.023-1.103]). A slowing of population growth started around 2000 (2000s: lambda = 1.030 [0.989-1.068]) and continued into the 2010s (lambda = 1.021 [0.985-1.057]), due primarily to reductions in survival of bears <2 years of age. These findings corroborate previous research that identified density-dependent effects as a likely cause. The IPM framework provided greater certainty and understanding regarding the dynamic demographic characteristics of the population and serves as a powerful monitoring tool for this long-lived species. Implementation of the IPM allows timely dissemination of demographic data to help inform adaptive management strategies and policy decisions necessary for the continued management and conservation of this population. This robust and flexible monitoring system allows scientists to investigate the effects of a changing ecosystem on population dynamics, incorporate new data sources and statistical models, and respond to changes in monitoring needs for the population. We highlight the efficacy of the IPM in estimating and tracking demographic parameters for a long-lived species, while accommodating shifts in monitoring techniques and data collections typical of long-term wildlife conservation programs worldwide.

Avoiding humans will be more difficult and energetically costly for animals as outdoor recreation increases and people venture farther into wildland areas that provide high-quality habitat for wildlife. Restricting human access can be an attractive management tool to mitigate effects of human recreation activities on wildlife; however, the efficacy of such measures is rarely assessed. In 1982, Yellowstone National Park identified areas important to grizzly bears (Ursus arctos) to help protect critical grizzly bear habitat and reduce the likelihood of human injuries by bears. Referred to as bear management areas (BMAs), human access is restricted in these areas for 2-8 months each year, with timing and type of restrictions varying by area. We examined 2 datasets to evaluate grizzly bear selection of BMAs and differences of bear density in BMAs and non-BMAs. First, we used 17 years of recent global positioning system telemetry data for grizzly bears to assess their selection of BMAs during periods when human access was allowed, and when access was restricted. We used step-selection functions to test the hypothesis that bears spend time in places that allow them to avoid people and select quality food sources. There was support that grizzly bears differentially select for BMAs regardless of whether human access was restricted at the time, compared with areas outside BMAs, and that selection changed with sex and season. Only males during the summer and hyperphagic seasons changed their selection of BMAs based on whether access restrictions were in place, and overall, male bears preferred unrestricted BMAs (BMAs without restrictions in place). Females preferentially selected BMAs regardless of whether the area had access restrictions in place only during the mating season. Individuals varied widely in their preference for BMAs and access restrictions. Bears likely choose to spend time in BMAs based on available food resources rather than restrictions to human access. Supporting this interpretation, our analyses indicated that a greater proportion of BMA in an area was associated with higher densities of grizzly bear. Thus, restrictions to human access likely help reduce the potential for human-bear interactions, accomplishing one of the original objectives for establishing the BMAs.

Understanding the density-dependent processes that drive population demography in a changing world is critical in ecology, yet measuring performance-density relationships in long-lived mammalian species demands long-term data, limiting scientists' ability to observe such mechanisms. We tested performance-density relationships for an opportunistic omnivore, grizzly bears (Ursus arctos, Linnaeus, 1758) in the Greater Yellowstone Ecosystem, with estimates of body composition (lean body mass and percent body fat) serving as indicators of individual performance over two decades (2000-2020) during which time pronounced environmental changes have occurred. Several high-calorie foods for grizzly bears have mostly declined in recent decades (e.g., whitebark pine [Pinus albicaulis, Engelm, 1863]), while increasing human impacts from recreation, development, and long-term shifts in temperatures and precipitation are altering the ecosystem. We hypothesized that individual lean body mass declines as population density increases (H1), and that this effect would be more pronounced among growing individuals (H2). We also hypothesized that omnivory helps grizzly bears buffer energy intake from changing foods, with body fat levels being independent from population density and environmental changes (H3). Our analyses showed that individual lean body mass was negatively related to population density, particularly among growing-age females, supporting H1 and partially H2. In contrast, population density or sex had little effect on body fat levels and rate of accumulation, indicating that sufficient food resources were available on the landscape to accommodate successful use of shifting food sources, supporting H3. Our results offer important insights into ecological feedback mechanisms driving individual performances within a population undergoing demographic and ecosystem-level changes. However, synergistic effects of continued climate change and increased human impacts could lead to more extreme changes in food availability and affect observed population resilience mechanisms. Our findings underscore the importance of long-term studies in protected areas when investigating complex ecological relationships in an increasingly anthropogenic world.

Conservation planning and decision-making can be enhanced by ecological models that reliably transfer to times and places beyond those where models were developed. Transferrable models can be especially helpful for species of conservation concern, such as grizzly bears (Ursus arctos). Currently, only four grizzly bear populations remain in the contiguous United States. We evaluated transferability of previously derived individual-based, integrated step selection functions (iSSFs) developed from GPS-collared grizzly bears in the Northern Continental Divide Ecosystem by applying them within the nearby Selkirk (SE), Cabinet-Yaak (CYE), and Greater Yellowstone Ecosystems (GYE). We simulated 100 replicates of 5000 steps for each iSSF in each ecosystem, summarized relative use into 10 equal-area classes for each sex, and overlaid GPS locations from bears in the SE, CYE, and GYE on resulting maps. Spearman rank correlations between numbers of locations and class rank were >= 0.96 within each study area, indicating models were highly predictive of grizzly bear space use in these nearby populations. Assessment of models using smaller subsets of data in space and time demonstrated generally high predictive accuracy for females. Although generally high across space and time, predictive accuracy for males was low within some watersheds and in summer within the SE and CYE, potentially due to seasonal effects, vegetation, and food assemblage differences. Altogether, these results demonstrated high transferability of our models to landscapes in the Northern Rocky Mountains, suggesting they may be used to evaluate habitat suitability and connectivity throughout the region to benefit conservation planning.

SUMMARYColor variation is a frequent evolutionary substrate for camouflage in small mammals but the underlying genetics and evolutionary forces that drive color variation in natural populations of large mammals are mostly unexplained. The American black bear, Ursus americanus, exhibits a range of colors including the cinnamon morph which has a similar color to the brown bear, U. arctos, and is found at high frequency in the American southwest. Reflectance and chemical melanin measurements showed little distinction between U. arctos and cinnamon U. americanus individuals. We used a genome-wide association for hair color as a quantitative trait in 151 U. americanus individuals and identified a single major locus (P < 10−13). Additional genomic and functional studies identified a missense alteration (R153C) in Tyrosinase-related protein 1 (TYRP1) that impaired protein localization and decreased pigment production. Population genetic analyses and demographic modeling indicated that the R153C variant arose 9.36kya in a southwestern population where it likely provided a selective advantage, spreading both northwards and eastwards by gene flow. A different TYRP1 allele, R114C, contributes to the characteristic brown color of U. arctos, but is not fixed across the range.HIGHLIGHTSThe cinnamon morph of American black bears and brown bears have different missense mutations in TYRP1 that account for their similar colorationTYRP1 variants in American black bears and brown bears are loss-of-function alleles associated with impaired protein localization to melanosomesIn American black bears, the variant causing the cinnamon morph arose 9,360 years ago in the western lineage where it provides an adaptive advantage, and has spread northwards and eastwards by migration

In the Greater Yellowstone Ecosystem, counts of female grizzly bears (Ursus arctos) with cubs-of-the-year (females with cubs) from systematic aerial surveys and opportunistic ground sightings are combined with demographic data to derive annual population estimates. We addressed 2 limitations to the monitoring approach. As part of a rule set, a conservative distance of >30 km currently is used as a threshold to assign sightings to unique females with cubs, resulting in underestimation bias. Using telemetry locations of females with cubs collected during 1997-2019, we created 1,000 data sets for each of 5 levels of simulated number of females with cubs, simulated sightings by selecting among these locations, and evaluated the classification performance of alternative distance criteria (1230 km). Under all scenarios, 12-16-km criteria maximized classification performance and minimized estimation bias; the 16-km criterion was optimal for current conditions and sampling efforts. Our second objective was to test generalized additive models (GAMs) as a flexible trend analysis technique. We simulated 1,000 time series for each of 10 scenarios (10, 15, and 20% decline over periods of 5, 10, and 15 yrs, plus stability), applied GAMs, and assessed metrics associated with the posterior distribution of the instantaneous rate of change. We detected declines among >99.6% of replicates under the 15 and 20% decline scenarios and in 84.7-94.7% of replicates under the 10% decline scenario. From decline onset to first detection, periods ranged from 3.7 (20% decline over 5 yrs) to 11.1 (10% decline over 15 yrs), with 3.9-8.8 years mean duration of detection events. The GAM approach allows detection of directional changes in population trend, including early warning metrics, and stabilization after such changes. Retrospective application of the 16-km distance criterion and GAMs resulted in higher population estimates and growth rates than are reported using current methods.

ABSTRACT Animal movements are major determinants of energy expenditure and ultimately the cost–benefit of landscape use. Thus, we sought to understand those costs and how grizzly bears (Ursus arctos) move in mountainous landscapes. We trained captive grizzly bears to walk on a horizontal treadmill and up and down 10% and 20% slopes. The cost of moving upslope increased linearly with speed and slope angle, and this was more costly than moving horizontally. The cost of downslope travel at slower speeds was greater than the cost of traveling horizontally but appeared to decrease at higher speeds. The most efficient walking speed that minimized cost per unit distance was 1.19±0.11 m s−1. However, grizzly bears fitted with GPS collars in the Greater Yellowstone Ecosystem moved at an average velocity of 0.61±0.28 m s−1 and preferred to travel on near-horizontal slopes at twice their occurrence. When traveling uphill or downhill, grizzly bears chose paths across all slopes that were ∼54% less steep and costly than the maximum available slope. The net costs (J kg−1 m−1) of moving horizontally and uphill were the same for grizzly bears, humans and digitigrade carnivores, but those costs were 46% higher than movement costs for ungulates. These movement costs and characteristics of landscape use determined using captive and wild grizzly bears were used to understand the strategies that grizzly bears use for preying on large ungulates and the similarities in travel between people and grizzly bears that might affect the risk of encountering each other on shared landscapes.

Meat, fruit, seeds and other high-energy bear foods are often highly localized and briefly available and understanding which factors influence bear consumption of these foods is a common focus of bear conservation and ecology. However, the most common bear foods, graminoids and forbs, are more widespread but of lower quality. We poorly understand how herbage consumption impacts bear physiology, such as endocrine system function that regulates homeostasis and stress responses. Here, we described bear diets with a novel approach, measuring the concentration of chlorophyll in bear scats (faecal chlorophyll) to index the proportion of the recent diet that was composed of leaves from graminoids and forbs. We measured faecal chlorophyll and faecal cortisol in 351 grizzly (Ursus arctos, n = 255) and black bear (Ursus americanus, n = 96) scats from Yellowstone National Park in 2008-2009. We compared models of faecal chlorophyll and faecal cortisol concentrations considering the effects of spatial, dietary, scat and bear-specific factors including species. Faecal chlorophyll levels were the strongest predictor of faecal cortisol in a manner that suggested an endocrine response to a low-energy diet. Both compounds were highest during the spring and early summer months, overlapping the breeding season when higher energy foods were less available. Effects of scat composition, scat weathering, bear age, bear sex, species and other factors that have previously been shown to influence faecal cortisol in bears were not important unless faecal chlorophyll was excluded from models. The top models of faecal chlorophyll suggested grazing was primarily influenced by spatial attributes, with greater grazing closer to recreational trails, implying that elevated cortisol with grazing could be a response to anthropogenic activity. Our results confirm that higher stress hormone concentrations correspond with lower quality diets in bears, particularly grazing, and that faecal chlorophyll shows promise as a metric for studying grazing behaviour and its consequences.

Heat dissipation limit theory posits that energy available for growth and reproduction in endotherms is limited by their ability to dissipate heat. In mammals, endogenous heat production increases markedly during gestation and lactation, and thus female mammals may be subject to greater thermal constraints on energy expenditure than males. Such constraints likely have important implications for behaviour and population performance in a warming climate. We used a mechanistic simulation model based on the first principles of heat and mass transfer to study thermal constraints on activity (both timing and intensity) of captive female grizzly bears Ursus arctos in current and future climate scenarios. We then quantified the relative importance of regulatory behaviours for maintaining heat balance using GPS telemetry locations of lactating versus non-lactating female bears from Yellowstone National Park, and assessed the degree to which costs of thermoregulation constrained the distribution of sampled bears in space and time. Lactating female bears benefitted considerably more from behavioural cooling mechanisms (e.g. partial submersion in cool water or bedding on cool substrate) than non-lactating females in our simulations; the availability of water for thermoregulation increased the number of hours during which lactating females could be active by up to 60% under current climatic conditions and by up to 43% in the future climate scenario. Moreover, even in the future climate scenario, lactating bears were able to achieve heat balance 24 hr/day by thermoregulating behaviourally when water was available to facilitate cooling. The most important predictor of female grizzly bear distribution in Yellowstone, regardless of reproductive status, was elevation. However, variables associated with the thermal environment were relatively more important for predicting the distribution of lactating than non-lactating female bears. Our results suggest that the costs of heat dissipation, which are modulated by climate, may impose constraints on the behaviour and energetics of large endotherms like grizzly bears, and that access to water for cooling will likely be an increasingly important driver of grizzly bear distribution in Yellowstone as the climate continues to warm. A free Plain Language Summary can be found within the Supporting Information of this article.

Animal movement is a complex phenomenon where individual movement patterns can be influenced by a variety of factors including the animal's current activity, available terrain and habitat, and locations of other animals. Motivated by modeling grizzly bear movement in the Greater Yellowstone Ecosystem, this article presents an agent-based model represented in a state-space framework for collective animal movement. The novel contribution of this work is a collective animal movement model that captures interactions between animals that can trigger changes in movement patterns, such as when a dominant grizzly bear may cause another subordinate bear to temporarily leave an area. The modeling framework enables learning different movement patterns through a state-space representation with particle-MCMC methods for fully Bayesian model fitting and the prediction of future animal movement behaviors.Supplementary materials accompanying this paper appear online.