Functional responses in polar bear habitat selection

Habitat selection may occur in situations in which animals experience a trade-off, e.g. between the use of habitats with abundant forage and the use of safer retreat habitats with little forage. Such trade-offs may yield relative habitat use conditional on the relative availability of the different habitat types, as proportional use of foraging habitat may exceed proportional availability when foraging habitat is scarce, but be less than availability when foraging habitat is abundant. Hence, trade-offs in habitat use may result in functional responses in habitat use (i.e. change in relative use with changing availability). We used logistic and log-linear models to model functional responses in female polar bear habitat use based on satellite telemetry data from two contiguous populations; one near shore inhabiting sea ice within fjords, and one inhabiting pelagic drift ice. Open ice, near the ice edge, is a highly dynamic habitat hypothesised to be important polar bear habitat due to high prey availability. In open ice-polar bears may experience a high energetic cost of movements and risk drifting away from the main ice field (i.e. trade off between feeding and energy saving or safety). If polar bears were constrained by ice dynamics we therefore predicted use of retreat habitats with greater ice coverage relative to habitats used for hunting. The polar bears demonstrated season and population specific functional responses in habitat use, likely reflecting seasonal and regional variation in use of retreat and foraging habitats. We suggest that in seasons with functional responses in habitat use, polar bear space use and population distribution may not be a mere reflection of prey availability but rather reflect the alternate allocation of time in hunting and retreat habitats.     

Habitat degradation affects the summer activity of polar bears

Understanding behavioral responses of species to environmental change is critical to forecasting population-level effects. Although climate change is significantly impacting species’ distributions, few studies have examined associated changes in behavior. Polar bear (Ursus maritimus) subpopulations have varied in their near-term responses to sea ice decline. We examined behavioral responses of two adjacent subpopulations to changes in habitat availability during the annual sea ice minimum using activity data. Location and activity sensor data collected from 1989 to 2014 for 202 adult female polar bears in the Southern Beaufort Sea (SB) and Chukchi Sea (CS) subpopulations were used to compare activity in three habitat types varying in prey availability: (1) land; (2) ice over shallow, biologically productive waters; and (3) ice over deeper, less productive waters. Bears varied activity across and within habitats with the highest activity at 50–75% sea ice concentration over shallow waters. On land, SB bears exhibited variable but relatively high activity associated with the use of subsistence-harvested bowhead whale carcasses, whereas CS bears exhibited low activity consistent with minimal feeding. Both subpopulations had fewer observations in their preferred shallow-water sea ice habitats in recent years, corresponding with declines in availability of this substrate. The substantially higher use of marginal habitats by SB bears is an additional mechanism potentially explaining why this subpopulation has experienced negative effects of sea ice loss compared to the still-productive CS subpopulation. Variability in activity among, and within, habitats suggests that bears alter their behavior in response to habitat conditions, presumably in an attempt to balance prey availability with energy costs.     

Are polar bear habitat resource selection functions developed from 1985–1995 data still useful?

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Polar bears in a warming climate

Polar bears (Ursus maritimus) live throughout the ice-coveredwaters of the circumpolar Arctic, particularly in near shoreannual ice over the continental shelf where biological productivityis highest. However, to a large degree under scenarios predictedby climate change models, these preferred sea ice habitats willbe substantially altered. Spatial and temporal sea ice changeswill lead to shifts in trophic interactions involving polarbears through reduced availability and abundance of their mainprey: seals. In the short term, climatic warming may improvebear and seal habitats in higher latitudes over continentalshelves if currently thick multiyear ice is replaced by annualice with more leads, making it more suitable for seals. A cascadeof impacts beginning with reduced sea ice will be manifestedin reduced adipose stores leading to lowered reproductive ratesbecause females will have less fat to invest in cubs duringthe winter fast. Non-pregnant bears may have to fast on landor offshore on the remaining multiyear ice through progressivelylonger periods of open water while they await freeze-up anda return to hunting seals. As sea ice thins, and becomes morefractured and labile, it is likely to move more in responseto winds and currents so that polar bears will need to walkor swim more and thus use greater amounts of energy to maintaincontact with the remaining preferred habitats. The effects ofclimate change are likely to show large geographic, temporaland even individual differences and be highly variable, makingit difficult to develop adequate monitoring and research programs.All ursids show behavioural plasticity but given the rapid paceof ecological change in the Arctic, the long generation time,and the highly specialised nature of polar bears, it is unlikelythat polar bears will survive as a species if the sea ice disappearscompletely as has been predicted by some.     

Seasonal matching of habitat quality and fitness in a migratory bird

When species occupy habitats that vary in quality, choice of habitat can be critical in determining individual fitness. In most migratory species, juveniles migrate independently of their parents and must therefore choose both breeding and winter habitats. Using a unique dataset of marked black-tailed godwits (Limosa limosa islandica) tracked throughout their migratory range, combined with analyses of stable carbon isotope ratios, we show that those individuals that occupy higher quality breeding sites also use higher quality winter sites. This seasonal matching can severely inflate inequalities in individual fitness. This population has expanded over the last century into poorer quality breeding and winter habitats and, across the whole population; individual birds tend to occupy either novel or traditional sites in both seasons. Winter and breeding season habitat selection are thus strongly linked throughout this population; these links have profound implications for a wide range of population and evolutionary processes. As adult godwits are highly philopatric, the initial choice of winter habitat by juveniles will be critical in determining future survival, timing of migration and breeding success.     

Space Use and Habitat Selection by Female Louisiana Black Bears in the Tensas River Basin of Louisiana

Abstract: Studies of space use and habitat selection of endangered species are useful for identifying factors that influence fitness of individuals and viability of populations. However, there is a lack of published information regarding these behaviors for the federally threatened Louisiana black bear (Ursus americanus luteolus). We documented space use and habitat selection for 28 female black bears in 2 subpopulations of the Tensas River Basin population in northeast Louisiana, USA. The Tensas subpopulation inhabits a relatively large (>300-km²) contiguous area of bottomland hardwood forest, whereas the Deltic subpopulation exists mainly in 2 small (<7-km²) forested patches surrounded by an agricultural matrix. Females on Deltic maintained smaller seasonal and annual home ranges than females on Tensas (all P < 0.04), except for females with cubs during spring. On Tensas, females with cubs maintained smaller home ranges than females without cubs during spring (P = 0.01), but we did not detect this difference on Deltic or in other seasons. Females on Tensas and Deltic exhibited differences in habitat selection when establishing home ranges and within home ranges (P < 0.001). Deltic females selected mature bottomland hardwood forests and avoided agricultural habitats at both spatial scales. Tensas females selected a mixture of swamps, mature and regenerating forests, and exhibited variation in selection across scale, season, and reproductive status. We suggest that differences in space use and habitat selection between Tensas and Deltic are at least partially due to habitat differences at the landscape (i.e., amount of forested habitat) and patch (i.e., food availability) scales. Our results contribute to the understanding of factors that influence space use and habitat selection by black bears and provide specific information on habitat types selected by Louisiana black bears to agencies involved in habitat protection and restoration for this threatened subspecies.     

Long-term density-dependent changes in habitat selection in red deer (Cervus elaphus)

Understanding how habitat selection changes with population density is a key concept in population regulation, community composition and managing impacts on biodiversity and ecosystem services. At low density, it is expected that individuals select habitats in terms of their preference, but as population density increases, the availability of resources per individual declines on preferred habitats, leading to competition which forces some individuals to exploit less preferred habitats. Using spatial information of Scottish red deer (Cervus elaphus) winter counts, carried out in 110 areas across Scotland between 1961 and 2004 (a total of 1,206,495 deer observations), we showed how winter habitat niche breadth in red deer has widened with increasing population density. Heather moorland and montane habitats were most and least preferred for deer, respectively. Increasing density favoured the selection of grassland, to the detriment of the selection of heather moorland. The selection of heather and grassland decreased when temperature increased, while the selection of montane and peatland habitats increased. These findings are important for understanding how habitat use, density and population are likely to be affected by weather, and allow us to predict habitat impacts by large mammal herbivory and climate.     

Spatial and temporal patterns of problem polar bears in Churchill,Manitoba

Human–bear interactions near the town of Churchill, Manitoba occur annually because the Western Hudson Bay polar bear population spends 4–5 months on-land each year when the sea ice melts completely. Significant changes have occurred in the Hudson Bay ecosystem and in the bear population as a result of climate warming; however, how these changes may have influenced human–bear interactions near Churchill is unclear. This study examined the temporal and spatial patterns of 1,487 problem bears captured in the Churchill area from 1970 to 2004. We also examined the relationship between problem bears and environmental variables as well as the Nunavut harvest. The number of individual problem bears caught near Churchill varied from 10 to 90 individuals per year and increased over time. Subadult males comprised 39%, subadult females 23%, adult males 18%, females with young 14%, and solitary females 6% of captures. Bears that became problem individuals were in closer proximity to the Churchill area. Nutritional stress and a northward shift in the distribution of the bears that spend the summer on-land in northeastern Manitoba may account for the increase in problem bear numbers. The date of sea ice freeze-up, which is getting progressively later, was the best predictor explaining the annual variation in the occurrence of problem bears. These results provide an understanding of how a warming climate may directly impact polar bear behaviour. This information may allow wildlife managers to predict relative levels of human–bear interactions and thereby implement effective management strategies to improve human safety and the conservation of polar bears.     

Sea ice resource selection models for polar bears in the Barents Sea subpopulation

The extent, thickness and age of Arctic sea ice has dramatically declined since the late 1990s, and these trends are predicted to continue. Exploring the habitat use of sea‐ice‐dependent species can help us understand which resources they use and how their distribution responds to a changing environment. The goal of this study was to develop predictive models of the habitat use of an Arctic apex predator. Polar bear Ursus maritimus habitat use in the Barents Sea subpopulation was modelled with seasonal resource selection functions (RSFs) using satellite‐linked telemetry data from 294 collars deployed on female polar bears between 1991 and 2015. Polar bears selected habitat in the Marginal Ice Zone, with a preference for intermediate sea ice concentrations (40–80%). They spent most time in areas with relatively short travel distances to 15 or 75% ice concentration, and during spring and autumn they exhibited a preference for sea ice areas over the continental shelf or over shallower bathymetry). Predictions of the distribution of polar bears in the Barents Sea area can be made for specific sea ice scenarios using these models. Two such predictive distribution maps based on the autumn seasonal model were made and validated against two independent polar bear survey datasets collected in August 2004 and August 2015. The distribution of optimal polar bear habitat has shifted strongly northwards in all seasons of the year during the 25 yr study period.     

A piecewise linear modeling approach for testing competing theories of habitat selection: an example with mule deer in northern winter ranges

Habitat selection fundamentally drives the distribution of organisms across landscapes; density-dependent habitat selection (DDHS) is considered a central component of ecological theories explaining habitat use and population regulation. A preponderance of DDHS theories is based on ideal distributions, such that organisms select habitat according to either the ideal free, despotic, or pre-emptive distributions. Models that can be used to simultaneously test competing DDHS theories are desirable to help improve our understanding of habitat selection. We developed hierarchical, piecewise linear models that allow for simultaneous testing of DDHS theories and accommodate densities from multiple habitats and regional populations, environmental covariates, and random effects. We demonstrate the use of these models with data on mule deer (Odocoileus hemionus) abundance and net energy costs in different snow depths within winter ranges of five regional populations in western Idaho, USA. Regional population density explained 40 % of the variation in population growth, and we found that deer were ideal free in winter ranges. Deer occupied habitats with lowest net energy costs at higher densities and at a higher rate than compared to habitats with intermediate and high energy costs. The proportion of a regional population in low energy cost habitat the previous year accounted for a significant amount of variation in population growth (17 %), demonstrating the importance of winter habitat selection in regulating deer populations. These linear models are most appropriate for empirical data collected from centralized habitat patches within the local range of a species where individuals are either year-round residents or migratory (but have already arrived from migration).     

High interindividual variability in habitat selection and functional habitat relationships in European nightjars over a period of habitat change

An animal's choice of foraging habitat reflects its response to environmental cues and is likely to vary among individuals in a population. Analyzing the magnitude of individual habitat selection can indicate how resilient populations may be to anthropogenic habitat change, where individually varying, broadly generalist populations have the potential to adjust their behavior. We collected GPS point data from 39 European nightjars (Caprimulgus europaeus) at a UK breeding site where restoration measures have altered large areas of habitat between breeding seasons. We calculated individual habitat selection over four breeding seasons to observe changes that might align with change in habitat. We also analyzed change in home range size in line with change in habitat availability, to examine functional relationships that can represent trade‐offs made by the birds related to performance of the habitat. Individual explained more of the variation in population habitat selection than year for most habitat types. Individuals differed in the magnitude of their selection for different habitat types, which created a generalist population composed of both generalist and specialist individuals. Selection also changed over time but only significantly for scrub habitat (60% decrease in selection over 4 years). Across the population, individual home range size was 2% smaller where availability of cleared habitat within the home range was greater, but size increased by 2% where the amount of open water was higher, indicating the presence of trade‐offs related to habitat availability. These results highlight that using individual resource selection and specialization measures, in conjunction with functional responses to change, can lead to better understanding of the needs of a population. Pooling specialist and generalist individuals for analysis could hide divergent responses to change and consequently obscure information that could be important in developing effective conservation strategies.     

Range contraction and increasing isolation of a polar bear subpopulation in an era of sea‐ice loss

Climate change is expected to result in range shifts and habitat fragmentation for many species. In the Arctic, loss of sea ice will reduce barriers to dispersal or eliminate movement corridors, resulting in increased connectivity or geographic isolation with sweeping implications for conservation. We used satellite telemetry, data from individually marked animals (research and harvest), and microsatellite genetic data to examine changes in geographic range, emigration, and interpopulation connectivity of the Baffin Bay (BB) polar bear (Ursus maritimus) subpopulation over a 25‐year period of sea‐ice loss. Satellite telemetry collected from n = 43 (1991–1995) and 38 (2009–2015) adult females revealed a significant contraction in subpopulation range size (95% bivariate normal kernel range) in most months and seasons, with the most marked reduction being a 70% decline in summer from 716,000 km² (SE 58,000) to 211,000 km² (SE 23,000) (p < .001). Between the 1990s and 2000s, there was a significant shift northward during the on‐ice seasons (2.6^° shift in winter median latitude, 1.1^° shift in spring median latitude) and a significant range contraction in the ice‐free summers. Bears in the 2000s were less likely to leave BB, with significant reductions in the numbers of bears moving into Davis Strait (DS) in winter and Lancaster Sound (LS) in summer. Harvest recoveries suggested both short and long‐term fidelity to BB remained high over both periods (83–99% of marked bears remained in BB). Genetic analyses using eight polymorphic microsatellites confirmed a previously documented differentiation between BB, DS, and LS; yet weakly differentiated BB from Kane Basin (KB) for the first time. Our results provide the first multiple lines of evidence for an increasingly geographically and functionally isolated subpopulation of polar bears in the context of long‐term sea‐ice loss. This may be indicative of future patterns for other polar bear subpopulations under climate change.     

Habitat‐mediated timing of migration in polar bears: an individual perspective

Migration phenology is largely determined by how animals respond to seasonal changes in environmental conditions. Our perception of the relationship between migratory behavior and environmental cues can vary depending on the spatial scale at which these interactions are measured. Understanding the behavioral mechanisms behind population‐scale movements requires knowledge of how individuals respond to local cues. We show how time‐to‐event models can be used to predict what factors are associated with the timing of an individual's migratory behavior using data from GPS collared polar bears (Ursus maritimus) that move seasonally between sea ice and terrestrial habitats. We found the concentration of sea ice that bears experience at a local level, along with the duration of exposure to these conditions, was most associated with individual migration timing. Our results corroborate studies that assume thresholds of >50% sea ice concentration are necessary for suitable polar bear habitat; however, continued periods (e.g., days to weeks) of exposure to suboptimal ice concentrations during seasonal melting were required before the proportion of bears migrating to land increased substantially. Time‐to‐event models are advantageous for examining individual movement patterns because they account for the idea that animals make decisions based on an accumulation of knowledge from the landscapes they move through and not simply the environment they are exposed to at the time of a decision. Understanding the migration behavior of polar bears moving between terrestrial and marine habitat, at multiple spatiotemporal scales, will be a major aspect of quantifying observed and potential demographic responses to climate‐induced environmental changes.     

Female polar bears, Ursus maritimus, on the Barents Sea drift ice: walking the treadmill

For animals in dynamic habitats, the contribution of passive (i.e. by wind or current) and active (movements by the animals themselves) displacement determines whether their space use reflects physical or adaptive behavioural processes. Polar bears in the Barents Sea undertake extensive annual migrations in a habitat that is highly dynamic because of continuous sea ice drift. Using combined information from satellite telemetry, satellite images and atmospheric pressure recordings, we estimated the contribution of sea ice drift and movements in the monthly net displacement of female polar bears. We found that movements, and thus behavioural processes, were dominant. Net displacement was directed northwards during summer ice retreat and southwards during winter ice advance. Conversely, movements were directed northwards counteracting a continuous southward drift. Acting as a treadmill, ice drift probably increased the energetic cost of migrations relative to that expected from observed net displacement distances; this suggests that pelagic and adjacent near-shore bears, on stable land-fast ice, have different energy costs. Little concordance between ice drift rates and net displacement and movement rates suggest that polar bears do not adjust their displacement relative to attractive areas with fixed locations, but rather adjust their movements to local habitat suitability. Furthermore, selective use of less dynamic drift ice when with cubs-of-the-year, and use of terrestrial denning areas, appear to be behavioural adaptations to the dynamics of the Barents Sea drift ice. Hence, understanding the behaviour and ecology of animals inhabiting dynamic habitats necessitates incorporation of both dynamic and static habitat variables. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Unpredictable selection in a structured population leads to local genetic differentiation in evolved reaction norms

Unpredictability during development of the optimum phenotype under future selection leads to a compromise reaction norm with a slope that is shallower than the slope of the optimum reaction norm. Unpredictability of selection can lead to an evolved curved reaction norm when genetic variation for curvature is available even if the optimum reaction norm is linear. This requires asymmetry in the frequency distribution of the habitats of selection; at small population size, stochasticity in the number of individuals per selection habitat is sufficient to generate such asymmetry. Unpredictability of selection in structured populations leads to local genetic differentiation of reaction norms. The mean habitat of a subpopulation is defined as the subpopulation's focal habitat. The evolved mean reaction norm of each subpopulation is anchored at the optimum genotypic value in its focal habitat. Linear reaction norms are parallel if the conditional distribution of adults around the focal habitats is the same for each subpopulation. Adult migration and absence of zygote dispersal represents the ultimate structured population, each habitat playing the role of focal habitat. Absence of zygote dispersal requires that the flow of individuals through the habitats is used instead of the habitats’ frequencies in the prediction of the evolved reaction norm. Adult migration in absence of zygote dispersal leads to an evolved pattern of locally differentiated reaction norms with optimum genotypic value anchored in the focal habitat and, for linear reaction norms, parallel slopes.     

Assessing year-round habitat use by migratory sea ducks in a multi-species context reveals seasonal variation in habitat selection and partitioning

Long-distance migration presents complex conservation challenges, and migratory species often experience shortfalls in conservation due to the difficulty of identifying important locations and resources throughout the annual cycle. In order to prioritize habitats for conservation of migratory wildlife, it is necessary to understand how habitat needs change throughout the annual cycle, as well as to identify key habitat sites and features that concentrate large numbers of individuals and species. Among long-distance migrants, sea ducks have particularly complex migratory patterns, which often include distinct post-breeding molt sites as well as breeding, staging and wintering locations. Using a large set of individual tracking data (n = 476 individuals) from five species of sea ducks in eastern North America, we evaluated multi-species habitat suitability and partitioning across the breeding, post-breeding migration and molt, wintering and pre-breeding migration seasons. During breeding, species generally occupied distinct habitat areas, with the highest levels of multi-species overlap occurring in the Barrenlands west of Hudson Bay. Species generally preferred flatter areas closer to lakes with lower maximum temperatures relative to average conditions, but varied in distance to shore, elevation and precipitation. During non-breeding, species overlapped extensively during winter but diverged during migration. All species preferred shallow-water, nearshore habitats with high productivity, but varied in their relationships to salinity, temperature and bottom slope. Sea ducks selected most strongly for preferred habitats during post-breeding migration, with high partitioning among species; however, both selection and partitioning were weaker during pre-breeding migration. The addition of tidal current velocity, aquatic vegetation presence and bottom substrate improved non-breeding habitat models where available. Our results highlight the utility of multi-species, annual-cycle habitat assessments in identifying key habitat features and periods of vulnerability in order to optimize conservation strategies for migratory wildlife.     

The association of Antarctic krill Euphausia superba with the under-ice habitat

The association of Antarctic krill Euphausia superba with the under-ice habitat was investigated in the Lazarev Sea (Southern Ocean) during austral summer, autumn and winter. Data were obtained using novel Surface and Under Ice Trawls (SUIT), which sampled the 0-2 m surface layer both under sea ice and in open water. Average surface layer densities ranged between 0.8 individuals m(-2) in summer and autumn, and 2.7 individuals m(-2) in winter. In summer, under-ice densities of Antarctic krill were significantly higher than in open waters. In autumn, the opposite pattern was observed. Under winter sea ice, densities were often low, but repeatedly far exceeded summer and autumn maxima. Statistical models showed that during summer high densities of Antarctic krill in the 0-2 m layer were associated with high ice coverage and shallow mixed layer depths, among other factors. In autumn and winter, density was related to hydrographical parameters. Average under-ice densities from the 0-2 m layer were higher than corresponding values from the 0-200 m layer collected with Rectangular Midwater Trawls (RMT) in summer. In winter, under-ice densities far surpassed maximum 0-200 m densities on several occasions. This indicates that the importance of the ice-water interface layer may be under-estimated by the pelagic nets and sonars commonly used to estimate the population size of Antarctic krill for management purposes, due to their limited ability to sample this habitat. Our results provide evidence for an almost year-round association of Antarctic krill with the under-ice habitat, hundreds of kilometres into the ice-covered area of the Lazarev Sea. Local concentrations of postlarval Antarctic krill under winter sea ice suggest that sea ice biota are important for their winter survival. These findings emphasise the susceptibility of an ecological key species to changing sea ice habitats, suggesting potential ramifications on Antarctic ecosystems induced by climate change.     

Variation in the response of an Arctic top predator experiencing habitat loss: feeding and reproductive ecology of two polar bear populations

Polar bears (Ursus maritimus) have experienced substantial changes in the seasonal availability of sea ice habitat in parts of their range, including the Beaufort, Chukchi, and Bering Seas. In this study, we compared the body size, condition, and recruitment of polar bears captured in the Chukchi and Bering Seas (CS) between two periods (1986–1994 and 2008–2011) when declines in sea ice habitat occurred. In addition, we compared metrics for the CS population 2008–2011 with those of the adjacent southern Beaufort Sea (SB) population where loss in sea ice habitat has been associated with declines in body condition, size, recruitment, and survival. We evaluated how variation in body condition and recruitment were related to feeding ecology. Comparing habitat conditions between populations, there were twice as many reduced ice days over continental shelf waters per year during 2008–2011 in the SB than in the CS. CS polar bears were larger and in better condition, and appeared to have higher reproduction than SB bears. Although SB and CS bears had similar diets, twice as many bears were fasting in spring in the SB than in the CS. Between 1986–1994 and 2008–2011, body size, condition, and recruitment indices in the CS were not reduced despite a 44‐day increase in the number of reduced ice days. Bears in the CS exhibited large body size, good body condition, and high indices of recruitment compared to most other populations measured to date. Higher biological productivity and prey availability in the CS relative to the SB, and a shorter recent history of reduced sea ice habitat, may explain the maintenance of condition and recruitment of CS bears. Geographic differences in the response of polar bears to climate change are relevant to range‐wide forecasts for this and other ice‐dependent species.     

Increased Land Use by Chukchi Sea Polar Bears in Relation to Changing Sea Ice Conditions

Recent observations suggest that polar bears (Ursus maritimus) are increasingly using land habitats in some parts of their range, where they have minimal access to their preferred prey, likely in response to loss of their sea ice habitat associated with climatic warming. We used location data from female polar bears fit with satellite radio collars to compare land use patterns in the Chukchi Sea between two periods (1986–1995 and 2008–2013) when substantial summer sea-ice loss occurred. In both time periods, polar bears predominantly occupied sea-ice, although land was used during the summer sea-ice retreat and during the winter for maternal denning. However, the proportion of bears on land for > 7 days between August and October increased between the two periods from 20.0% to 38.9%, and the average duration on land increased by 30 days. The majority of bears that used land in the summer and for denning came to Wrangel and Herald Islands (Russia), highlighting the importance of these northernmost land habitats to Chukchi Sea polar bears. Where bears summered and denned, and how long they spent there, was related to the timing and duration of sea ice retreat. Our results are consistent with other studies supporting increased land use as a common response of polar bears to sea-ice loss. Implications of increased land use for Chukchi Sea polar bears are unclear, because a recent study observed no change in body condition or reproductive indices between the two periods considered here. This result suggests that the ecology of this region may provide a degree of resilience to sea ice loss. However, projections of continued sea ice loss suggest that polar bears in the Chukchi Sea and other parts of the Arctic may increasingly use land habitats in the future, which has the potential to increase nutritional stress and human-polar bear interactions.