Migration conveys many benefits to species, ecosystems, and people but relies upon connected landscapes. Anthropogenic development can present barriers for migrants, but many barriers are semi-permeable, allowing unhindered or delayed passage. We used a modified version of the Barrier Behavior Analysis (BaBA) to investigate seasonal movement responses to five roads in northwestern Alaska by adult female Western Arctic Herd caribou (Rangifer tarandus) from 2009 to 2024. Our analyses revealed some altered movement in response to all focal roads. We found that the roads were semi-permeable barriers to movement, with altered behaviors including bouncing away, moving back-and-forth, and tracing along roads. Overall, 63.1% of collared animals encountered (entered a road-specific buffer) at least one focal road. Of these, 61.5% displayed altered movements. At the scale of individual encounters with roads, we found altered movement in 27.1% of road encounters. Most encounters occurred during fall migration and caribou with altered behavior spent an average of 9.4 days longer near focal roads than those with unaltered movement. Altered movements were balanced among the behavioral responses. Most altered movements occurred near the Red Dog mining road (60.3%) or during fall migration (51.9%) but lasted longest during winter (17.2 days on average). We confirm prior findings of altered fall movements near the Red Dog road and demonstrate that movement behavior is also altered around other roads and in other seasons. Nonetheless, many collared caribou did not display altered movements in response to roads, emphasizing the need for further research to understand the mechanistic drivers of caribou movement responses. Given increasing pressures for infrastructure development and global challenges facing migratory species, it is critical to identify mitigation measures and inform management decisions seeking to balance responsible development with conservation of natural systems, including migratory species and the people that rely upon them.

Understanding animal behavior at the population level can be challenging, especially in the presence of intraspecific variation in behavioral tactics. Individuals within a population often vary with respect to resource exploitation and use, which may be associated with individual states (e.g., male or female) or extrinsic variation (e.g., temporal variation in food). Explicitly accounting for interindividual variation can aid ecological insights, especially for species that exhibit high behavioral flexibility. Here, we evaluated how seasonal fluctuations in resources influenced individual-level habitat use and selection of Rocky Mountain red fox (Vulpes vulpes macroura) within Grand Teton National Park, Wyoming. Rocky Mountain red fox is a high-elevation subspecies of the common red fox (Vulpes vulpes), and native to North America. From 2016 to 2021, we captured and GPS-collared 18 individuals (11 males, 7 females) and evaluated spatiotemporal shifts in home range characteristics and habitat selection. We observed high individual variation in home range size, with generally larger home ranges in the winter months (x$$ \overline{x} $$ = 72.92 km2 [minimum convex polygon-MCP], 83.65 km2 [Kernel], 27.20 km2 [Local Convex Hull]) than in the summer (x$$ \overline{x} $$ = 22.23 km2 [MCP], 23.01 km2 [Kernel], 11.11 km2 [Local Convex Hull]). Similarly, we observed substantial individual variation in habitat selection across environmental gradients. Some foxes altered their selection for habitat types between summer and winter indicating behavioral plasticity with respect to seasonal resources. Distance to human features was the primary driver for habitat selection for both seasons across foxes, and on average, foxes had stronger selection for human features in the summer. These findings might indicate some foxes are concentrating on the temporal and spatial resource pulse of anthropogenic food, while others exhibited more natural tactics. Our work advanced the spatial ecology of Rocky Mountain red fox, identified variation in space use tactics associated with humans and natural resources among red foxes in a heavily recreated area, and demonstrated how human activity can influence the spatial behavior of a carnivore within a national park.

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.

Denning is a critical behavioral adaptation for brown bears Ursus arctos to cope with winter, a period of extended resource scarcity. Bears reduce their body temperature, heart rate, and metabolism during this time to minimize energy expenditures. The Arctic has among the most pronounced and longest period of resource scarcity. Thus, we predicted bears in the region would respond by having among the longest recorded denning periods. We used GPS data from brown bears to determine the den entry, den exit, and denning duration for a population living primarily above the Arctic Circle. On average, brown bears in the region denned for 206 days, the longest duration reported using GPS data of which we are aware. The longest denning duration for any individual bear was a remarkable 233 days (64% of the year), which is near the theoretical maximum of 241 days. We found that food availability in fall delayed den entrance, with bears that appeared to consume more salmon entering their dens later. Bears showed greater synchrony in den exiting than den entrance, and female bears with cubs exited their dens more than a week after other bears. Later snow melt out in spring was also associated with later den exits. Climate change has the potential to affect the denning ecology of Arctic brown bears by altering the availability of food, ambient temperature, and precipitation, all of which can alter the costs and benefits of hibernation for brown bears.

Among polar bears (Ursus maritimus), only parturient females den for extended periods, emerging from maternal dens in spring after having substantially depleted their energy reserves during a fast that can exceed 8 months. Although den emergence coincides with a period of increasing prey availability, polar bears typically do not depart immediately to hunt, but instead remain at the den for up to a month. This delay suggests that there are likely adaptive advantages to remaining at the den between emergence and departure, but the influence of the timing and duration of this post-emergence period on cub survival has not been evaluated previously. We used temperature and location data from 70 denning bears collared within the Southern Beaufort Sea and Chukchi Sea subpopulations to estimate the phenology of the post-emergence period. We evaluated the influence of various spatial and temporal features on duration of the post-emergence period and evaluated the potential influence of post-emergence duration on litter survival early in the spring following denning. For dens that likely contained viable cubs at emergence (n = 56), mean den emergence occurred on 16 March (SE = 1.4 days) and mean departure on 24 March (SE = 1.6 days), with dates typically occurring later in the Chukchi Sea relative to Southern Beaufort Sea and on land relative to sea ice. Mean duration of the post-emergence period was 7.9 days (SE = 1.4) for bears that were observed with cubs later in the spring, which was over 4 times longer than duration of those observed without cubs (1.9 days). Litter survival in the spring following denning (n = 31 dens) increased from 0.5 to 0.9 when duration of the post-emergence period increased by ~4 days and other variables were held at mean values. Our limited sample size and inability to verify cub presence at emergence suggests that future research is merited to improve our understanding of this relationship. Nonetheless, our results highlight the importance of the post-emergence period in contributing to reproductive success and can assist managers in developing conservation and mitigation strategies in denning areas, which will be increasingly important as human activities expand in the Arctic.

Mineral requirements are poorly described for most wildlife. Consequently, the role of forage minerals in movement and productivity are poorly understood for sedentary and migratory ungulates, such as reindeer and caribou (Rangifer tarandus). We applied estimates of maintenance, lactation, body mass change, and antler growth to production curves (body mass, daily intake, and milk yield) for female caribou to calculate their mineral requirements over summer. The total requirements (mg or g·d−1) were divided by the daily intake (kg·d−1) to estimate the minimum concentration of minerals required in the diet (mg or g·kg−1) to balance demand. The daily requirements (mg·d−1) of all minerals increased from parturition to the end of summer. The minimum dietary concentrations (mg·kg−1) of macro-minerals (Ca, P, Mg, Na, K) declined as food intake (kg·d−1) increased over summer. The minimum dietary concentrations (Fe, Mn, Cu, Zn) were heavily influenced by body mass gain, which increased through late lactation even though food intakes rose. Our modeling framework can be applied to other wild ungulates to assess the impacts of changing forage phenology, plant community compositions, or environmental disturbances on movement and productivity.

Providing novel food subsidies to wildlife can influence behavior, resource use, disease dynamics, and food web interactions. Studies examining impacts of anthropogenic food have primarily occurred in urban landscapes where anthropogenic resources are consistently accessible. National parks, however, experience similar issues regarding wildlife exploiting anthropogenic food, but access to anthropogenic food is contingent on human visitation, which can fluctuate dramatically between seasons. We evaluated how fluctuating availability of anthropogenic food and demographic factors influenced the isotopic niche of Rocky Mountain Red Fox (Vulpes vulpes macroura) at the population and individual level in Grand Teton National Park, Wyoming. From 2016 to 2021, we assessed diet-related questions by collecting 55 whisker samples and 52 hair samples from 40 unique individuals and analyzed them using stable isotope analysis (delta 13C and delta 15N). We discovered that Rocky Mountain red foxes have a broad isotopic niche, with seasonal differences in their consumption of anthropogenic food. Contrary to our expectations, foxes demonstrated increased reliance on anthropogenic food (higher delta 13C values) during winter months, whereas natural food consumption or dietary mixing was more prominent in summer months. Natural food scarcity could be driving foxes to seek out more anthropogenic food in the winter, whereas the increased abundance of natural food items in the summer could result in a lower need for foxes to supplement their diet with anthropogenic food. Adults had consistently higher delta 13C values relative to juveniles, indicating either intraspecific variation in boldness or dominance leading to increased exploitation of anthropogenic food, which could be a learned behavior coinciding with habituation. Finally, we observed that increasing densities of human features within the home range of a fox was a strong predictor of anthropogenic food reliance at the individual level. Our work provided a deeper understanding of the trophic ecology of Rocky Mountain Red Fox in relation to human activity in a nature preserve, and equipped managers with insight into potential patterns associated with food-conditioned foxes. How does human food influence carnivore feeding in a national park? We investigated how human food influenced the trophic ecology of the Rocky Mountain Red Fox within Grand Teton National Park. Spatial access was a driver of Red Fox consumption of human food, but consumption varied by season and age.

Contamination with arsenic (As), cadmium (Cd), mercury (Hg) and lead (Pb) is a global concern impairing resilience of organisms and ecosystems. Proximity to emission sources increases exposure risk but remoteness does not alleviate it. These toxic elements are transported in atmospheric and oceanic pathways and accumulate in organisms. Mercury accumulates in higher trophic levels. Brown bears (Ursus arctos), which often live in remote areas, are long-lived omnivores, feeding on salmon (Oncorhynchus spp.) and berries (Vaccinium spp.), resources also consumed by humans. We measured blood concentrations of As, Cd, Hg and Pb in bears (n = 72) four years and older in Scandinavia and three national parks in Alaska, USA (Lake Clark, Katmai and Gates of the Arctic) using high-resolution, inductively-coupled plasma sector field mass spectrometry. Age and sex of the bears, as well as the typical population level diet was associated with blood element concentrations using generalized linear regression models. Alaskan bears consuming salmon had higher Hg blood concentrations compared to Scandinavian bears feeding on berries, ants (Formica spp.) and moose (Alces). Cadmium and Pb blood concentrations were higher in Scandinavian bears than in Alaskan bears. Bears using marine food sources, in addition to salmon in Katmai, had higher As blood concentrations than bears in Scandinavia. Blood concentrations of Cd and Pb, as well as for As in female bears increased with age. Arsenic in males and Hg concentrations decreased with age. We detected elevated levels of toxic elements in bears from landscapes that are among the most pristine on the planet. Sources are unknown but anthropogenic emissions are most likely involved. All study areas face upcoming change: Increasing tourism and mining in Alaska and more intensive forestry in Scandinavia, combined with global climate change in both regions. Baseline contaminant concentrations as presented here are important knowledge in our changing world.

Large carnivores are recovering in many landscapes where the human footprint is simultaneously growing. When carnivores encounter humans, the way they behave often changes, which may subsequently influence how they affect their prey. However, little research investigates the behavioural mechanisms underpinning carnivore response to humans. As a result, it is not clear how predator-prey interactions and their associated ecosystem processes will play out in the human-dominated areas into which carnivore populations are increasingly expanding. We hypothesized that humans would reduce predation risk for prey by disturbing carnivores or threatening their survival. Alternatively, or additionally, we hypothesized that humans would increase predation risk by providing forage resources that congregate herbivorous prey in predictable places and times. Using grey wolves Canis lupus in Jackson Hole, Wyoming, USA as a study species, we investigated 170 kill sites across a spectrum of human influences ranging from heavily restricted human activities on protected federal lands to largely unregulated activities on private lands. Then, we used conditional logistic regression to quantify how the probability of predation changed across varied types and amounts of human influences, while controlling for environmental characteristics and prey availability. Wolves primarily made kills in environmental terrain traps and where prey availability was high, but predation risk was significantly better explained with the inclusion of human influences than by environmental characteristics alone. Different human influences had different, and even converse, effects on the risk of wolf predation. For example, where prey were readily available, wolves preferentially killed animals far from motorized roads but close to unpaved trails. However, wolves responded less strongly to humans, if at all, where prey were scarce, suggesting they prioritized acquiring prey over avoiding human interactions. Overall, our work reveals that the effects of large carnivores on prey populations can vary considerably among different types of human influences, yet carnivores may not appreciably alter predatory behaviour in response to humans if prey are difficult to obtain. These results shed new light on the drivers of large carnivore behaviour in anthropogenic areas while improving understanding of predator-prey dynamics in and around the wildland-urban interface.

Giardia and Cryptosporidium are zoonotic protozoan parasites that can infect humans and other taxa, including wildlife, often causing gastrointestinal illness. Both have been identified as One Health priorities in the Arctic, where climate change is expected to influence the distribution of many wildlife and zoonotic diseases, but little is known about their prevalence in local wildlife. To help fill information gaps, we collected fecal samples from four wildlife species that occur seasonally on the northern Alaska coastline or in nearshore marine waters-Arctic fox (Vulpes lagopus), polar bear (Ursus maritimus), Pacific walrus (Odobenus rosmarus divergens), and caribou (Rangifer tarandus)-and used immunofluorescence assays to screen for Giardia cysts and Cryptosporidium oocysts. We detected Giardia cysts in 18.3% and Cryptosporidium oocysts in 16.5% of Arctic foxes (n = 109), suggesting that foxes may be potentially important hosts in this region. We also detected Giardia cysts in a single polar bear (12.5%; n = 8), which to our knowledge represents the first such report for this species. Neither parasite was detected in walruses or caribou.

To assess infection with or exposure to endo- and ectoparasites in Alaska brown bears (Ursus arctos), blood and fecal samples were collected during 2013-17 from five locations: Gates of the Arctic National Park and Preserve; Katmai National Park; Lake Clark National Park and Preserve; Yakutat Forelands; and Kodiak Island. Standard fecal centrifugal flotation was used to screen for gastrointestinal parasites, molecular techniques were used to test blood for the presence of Bartonella and Babesia spp., and an ELISA was used to detect antibodies reactive to Sarcoptes scabiei, a species of mite recently associated with mange in American black bears (Ursus americanus). From fecal flotations (n=160), we identified the following helminth eggs: Uncinaria sp. (n=16, 10.0%), Baylisascaris sp. (n=5, 3.1%), Dibothriocephalus sp. (n=2, 1.2%), and taeniid-type eggs (n=1, 0.6%). Molecular screening for intraerythrocytic parasites (Babesia spp.) and intracellular bacteria (Bartonella spp.) was negative for all bears tested. We detected antibodies to S. scabiei in six of 59 (10.2%) individuals. The relatively low level of parasite detection in this study meets expectations for brown bear populations living in large, relatively undisturbed habitats near the northern edge of the range. These results provide a contemporary understanding of parasites in Alaska brown bears and establish baseline levels of parasite presence to monitor for changes over time and relative to ecologic alterations.

Investigators have speculated that the climate-driven “greening of the Arctic” may benefit barren-ground caribou populations, but paradoxically many populations have declined in recent years. This pattern has raised concerns about the influence of summer habitat conditions on caribou demographic rates, and how populations may be impacted in the future. The short Arctic summer provides caribou with important forage resources but is also the time they are exposed to intense harassment by insects, factors which are both being altered by longer, warmer growing seasons. To better understand the effects of summer forage and insect activity on Arctic caribou demographic rates, we investigated the influence of estimated forage biomass, digestible energy (DE), digestible nitrogen (DN), and mosquito activity on the reproductive success and survival of adult females in the Central Arctic Herd on the North Slope of Alaska. We tested the hypotheses that greater early summer DN would increase subsequent reproduction (parturition and late June calving success) while greater biomass and DE would increase adult survival (September–May), and that elevated mosquito activity would reduce both demographic rates. Because the period when abundant forage DN is limited and overlaps with the period of mosquito harassment, we also expected years with low DN and high harassment to synergistically reduce caribou reproductive success. Examining these relationships at the individual-level, using GPS-collared females, and at the population-level, using long-term monitoring data, we generally found support for our expectations. Greater early summer DN was associated with increased subsequent calving success, while greater summer biomass was associated with increased adult survival. Mosquito activity was associated with reductions in adult female parturition, late June calving success, and survival, and in years with low DN, had compounding effects on subsequent late June calving success. Our findings indicate that summer nutrition and mosquito activity collectively influence the demographic rates of Arctic caribou, and may impact the dynamics of populations in the future under changing environmental conditions.

Gut microbiomes (GMBs), complex communities of microorganisms inhabiting the gastrointestinal tracts of their hosts, perform countless micro-ecosystem services such as facilitating energy uptake and modulating immune responses. While scientists increasingly recognize the role GMBs play in host health, the role of GMBs in wildlife ecology and conservation has yet to be realized fully. Here, we use brown bears (Ursus arctos) as an ecological model to (1) characterize GMB community composition associated with location, season, and reproductive condition of a large omnivore; (2) investigate how both extrinsic and intrinsic factors influence GMB community membership and structure; and (3) quantify differences in GMB communities among different locations, seasons, sex, and reproductive conditions. To achieve these aims, we subsampled brown bear fecal samples collected during United States National Park Service research activities at three National Parks and Preserves (Katmai, Lake Clark, and Gates of the Arctic) and extracted microbial DNA for 16S rRNA amplicon sequencing and microbial taxonomic classification. We analyzed GMB communities using alpha and beta diversity indices, subsequently using linear mixed models to examine relationships between alpha diversity and extrinsic and intrinsic factors. Katmai brown bears hosted the greatest alpha diversity, whereas Gates brown bears hosted the least alpha diversity. Our results indicate that location and diet drive GMB variation, with bears hosting less phylogenetic diversity as park distance inland increases. Monitoring brown bear GMBs could enable managers to quickly detect and assess the impact of environmental perturbations on brown bear health. By integrating macro and micro-ecological perspectives we aim to inform local and landscape-level management decisions to promote long-term brown bear conservation and management.

Abstract: Home range size is a basic ecological index related to individual's realized niche. Its size can be influenced by body size, sex, maternal status, population density, habitat productivity, spatiotemporal variation of resources, climate, predation risk, and disturbance. Home range estimation can also be greatly affected by methodology and sampling regime. We used Global Positioning System collar data to assess what factors influenced the size of annual home ranges (space use during a single active season) of 28 female and 8 male brown bears (Ursus arctos) that denned in the Brooks Range of northcentral Alaska, USA, from 2014 to 2017. We used 2 methods to estimate annual home ranges, the Kernel Density Estimate (KDE) and the dynamic Brownian Bridge Movement Model (dBBMM). Contrary to expectations, we did not find that larger bodied bears of the same sex had larger annual home ranges. Annual home ranges of male bears (mean [standard deviation]; 504 [312] km2 and 3,886 [4,279] km2, using dBBMM and KDE, respectively) were 3.7–9.4 times larger than that of females (135 [86] km2 and 411 [738] km2, respectively). We found that greater chum salmon (Oncorhynchus keta) consumption was associated with larger annual home ranges for both sexes. In contrast, coastal brown bear populations that consume high levels of salmon often have small annual home ranges. We suggest that the relatively long distance (up to 100 km) between salmon streams and another key resource, denning habitat, is a reason for the positive association between salmon consumption and annual home range size. Although age was not in our top model for annual home range size, younger bears tended to have larger annual home ranges. We documented the fact that individuals of both sexes had the largest annual home ranges of any we could find for brown bears worldwide, using a traditional measure of space use (KDE). However, very large annual home ranges were associated with nonlocalized movements and the alternative method (dBBMM) to delineate these ranges provided more realistic range estimates. We discuss options and limitations of estimating space use and recommend caution when comparing space use between studies. With large-scale industrial infrastructure approved for development in this previously undeveloped region, the size and drivers of bear annual home ranges have numerous management implications. Brown bears with large annual home ranges in northcentral Alaska, where primary productivity is relatively low and denning habitat often far from salmon-bearing streams, are likely to move outside conservation units and encounter more risks as they interact with human infrastructure.

The internal mechanisms responsible for modulating physiological condition, particularly those performed by the gut microbiome (GMB), remain under-explored in wildlife. However, as latitudinal and seasonal shifts in resource availability occur, the myriad micro-ecosystem services facilitated by the GMB may be especially important to wildlife health and resilience. Here, we use brown bears ( Ursus arctos ) as an ecological model to quantify the relationship between wildlife body condition metrics that are commonly used to assess individual and population-level health and GMB community composition and structure. To achieve these aims, we subsampled brown bear fecal samples collected during United States National Park Service research activities at three National Parks and Preserves (Katmai, Lake Clark, and Gates of the Arctic) and extracted microbial DNA for 16S rRNA amplicon sequencing and microbial taxonomic classification. We analyzed GMB communities using alpha diversity indices, subsequently using Spearman’s correlation analysis to examine relationships between alpha diversity and brown bear health metrics. We found no differences in GMB composition among bears with differing body conditions, nor any correlations between alpha diversity and body condition. Our results indicate that GMB composition reflects diverse foraging strategies while allowing brown bears to achieve similar body condition outcomes.

Abstract Spatiotemporal variation in forage is a primary driver of ungulate behavior, yet little is known about the nutritional components they select, and how selection varies across the growing season with changes in forage quality and quantity. We addressed these uncertainties in barren‐ground caribou (Rangifer tarandus), which experience their most important foraging opportunities during the short Arctic summer. Recent declines in Arctic caribou populations have raised concerns about the influence of climate change on summer foraging opportunities, given shifting vegetation conditions and insect harassment, and their potential effects on caribou body condition and demography. We examined Arctic caribou selection of summer forage by pairing locations from females in the Central Arctic Herd of Alaska with spatiotemporal predictions of biomass, digestible nitrogen (DN), and digestible energy (DE). We then assessed selection for these nutritional components across the growing season at landscape and patch scales, and determined whether foraging opportunities were constrained by insect harassment. During early summer, at the landscape scale, caribou selected for intermediate biomass and high DN and DE, following expectations of the forage maturation hypothesis. At the patch scale, however, caribou selected for high values of all forage components, particularly DN, suggesting that protein may be limiting. During late summer, after DN declined below the threshold for protein gain, caribou exhibited a switch at both spatial scales, selecting for higher biomass, likely enabling mass and fat deposition. Mosquito activity strongly altered caribou selection of forage and increased their movement rates, while oestrid fly activity had little influence. Our results demonstrate that early and late summer periods afford Arctic caribou distinct foraging opportunities, as they prioritize quality earlier in the summer and quantity later. Climate change may further constrain caribou access to DN as earlier, warmer Arctic summers may be associated with reduced DN and increased mosquito harassment.