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The Arctic Ocean is undergoing rapid transformation toward a seasonally ice‐free ecosystem. As ice‐adapted apex predators, polar bears (Ursus maritimus) are challenged to cope with ongoing habitat degradation and changes in their prey base driven by food‐web response to climate warming. Knowledge of polar bear response to environmental change is necessary to understand ecosystem dynamics and inform conservation decisions. In the southern Beaufort Sea (SBS) of Alaska and western Canada, sea ice extent has declined since satellite observations began in 1979 and available evidence suggests that the carrying capacity of the SBS for polar bears has trended lower for nearly two decades. In this study, we investigated the population dynamics of polar bears in Alaska's SBS from 2001 to 2016 using a multistate Cormack–Jolly–Seber mark–recapture model. States were defined as geographic regions, and we used location data from mark–recapture observations and satellite‐telemetered bears to model transitions between states and thereby explain heterogeneity in recapture probabilities. Our results corroborate prior findings that the SBS subpopulation experienced low survival from 2003 to 2006. Survival improved modestly from 2006 to 2008 and afterward rebounded to comparatively high levels for the remainder of the study, except in 2012. Abundance moved in concert with survival throughout the study period, declining substantially from 2003 and 2006 and afterward fluctuating with lower variation around an average of 565 bears (95% Bayesian credible interval [340, 920]) through 2015. Even though abundance was comparatively stable and without sustained trend from 2006 to 2015, polar bears in the Alaska SBS were less abundant over that period than at any time since passage of the U.S. Marine Mammal Protection Act. The potential for recovery is likely limited by the degree of habitat degradation the subpopulation has experienced, and future reductions in carrying capacity are expected given current projections for continued climate warming., As ice‐adapted apex predators, polar bears (Ursus maritimus) are challenged to cope with the ongoing habitat degradation and changes in their prey base driven by food web response to climate warming. We investigated the population dynamics of polar bears in Alaska's southern Beaufort Sea (SBS) using a multistate CJS mark–recapture model. Abundance declined during several consecutive years of poor survival and afterward fluctuated in concert with survival but without sustained trend, averaging 565 bears from 2006 to 2015, the lowest level since passage of the US Marine Mammal Protection Act.

The analysis of animal tracking data provides important scientific understanding and discovery in ecology. Observations of animal trajectories using telemetry devices provide researchers with information about the way animals interact with their environment and each other. For many species, specific geographical features in the landscape can have a strong effect on behavior. Such features may correspond to a single point (eg, dens or kill sites), or to higher dimensional subspaces (eg, rivers or lakes). Features may be relatively static in time (eg, coastlines or home-range centers), or may be dynamic (eg, sea ice extent or areas of high-quality forage for herbivores). We introduce a novel model for animal movement that incorporates active selection for dynamic features in a landscape. Our approach is motivated by the study of polar bear (Ursus maritimus) movement. During the sea ice melt season, polar bears spend much of their time on sea ice above shallow, biologically productive water where they hunt seals. The changing distribution and characteristics of sea ice throughout the year mean that the location of valuable habitat is constantly shifting. We develop a model for the movement of polar bears that accounts for the effect of this important landscape feature. We introduce a two-stage procedure for approximate Bayesian inference that allows us to analyze over 300 000 observed locations of 186 polar bears from 2012 to 2016. We use our model to estimate a spatial boundary of interest to wildlife managers that separates two subpopulations of polar bears from the Beaufort and Chukchi seas.

Publisher Summary The polar bear is, on average, the largest of the eight species of bears found worldwide and is completely white. Ecologically, it is the only marine bear. Typically, the body is stocky, lacks a shoulder hump, and has a longer neck in relation to the rest of the body than other ursids. Polar bears are thought to have originated from brown bears, 1.3 million years ago. Polar bears and brown bears are capable of interbreeding in zoos and the young are fertile. Although in the wild, there is little overlap in the habitats during the breeding season, there has been one confirmed. Polar bears are distributed in ice-covered waters throughout the circumpolar Arctic. Although small numbers of animals may be found in the permanent multiyear pack ice of the central arctic basin, their preferred habitat is the annual ice over continental shelves of the continents and islands around the coastline of the Arctic Ocean where overall biological productivity and densities of ringed seals, their primary prey, are greatest. Regional concentrations and seasonal movement patterns of polar bears are influenced primarily by the type and distribution of sea ice and by the density and distribution of seals. Polar bears that have continuous access to sea ice continue to hunt throughout the year. Their hunting methods and rates of success change with the seasons and vary in different areas. The most common method of hunting is to lie beside a breathing hole waiting for a seal to surface and breathe. Polar bears have a remarkable ability to store large amounts of fat during periods when prey, mainly seals, are available and then fast for protracted periods when food is not available.

Divergent movement strategies have enabled wildlife populations to adapt to environmental change. In recent decades, the Southern Beaufort Sea subpopulation of polar bears (Ursus maritimus) has developed a divergent movement strategy in response to diminishing sea ice where the majority of the subpopulation (73-85%) stays on the sea ice in summer and the remaining bears move to land. Although declines in sea ice are generally considered a challenge to energy balance in polar bears residing in some regions of the Arctic, little quantitative data exists concerning the seasonal energy expenditures of this apex marine carnivore. We used GPS satellite collars with tri-axial accelerometers and conductivity sensors to measure the location, behavior, and energy expenditure of five adult female polar bears in the southern Beaufort Sea across seasons of sea ice breakup and minimum extent. Using a Bayesian mixed-effects model, we found that energy expenditure was influenced by month, ocean depth, and habitat type (sea ice or land). Total energy expenditure from May through September ranged from 37.7 to 47.2 mJ/kg for individual bears. Bears that moved to land expended 7% more energy on average from May through September than bears that remained on the receding sea ice. In August, when bears were moving from the sea ice to land or moving north with the receding pack ice, bears that moved to land spent 7% more time swimming and expended 22% more energy. This means the immediate cost of moving to land exceeded the cost of remaining on the receding summer pack ice. These findings suggest a physiological reason why the majority of the Southern Beaufort Sea subpopulation continues to inhabit a diminishing summer ice platform. However, bears that moved to land spent 29% more time in preferred hunting habitats over the continental shelf than bears that remained on the sea ice. Bears on land also had access to subsistence-harvested bowhead whale carcasses. Hence, our findings indicate there may be a greater overall energetic benefit to move to land in this region, which suggests that the use of the diminishing summer sea ice may be functioning as an ecological trap.

A demanding lifestyle Polar bears appear to be well adapted to the extreme conditions of their Arctic habitat. Pagano et al. , however, show that the energy balance in this harsh environment is narrower than we might expect (see the Perspective by Whiteman). They monitored the behavior and metabolic rates of nine free-ranging polar bears over 2 years. They found that high energy demands required consumption of high-fat prey, such as seals, which are easy to come by on sea ice but nearly unavailable in ice-free conditions. Thus, as sea ice becomes increasingly short-lived annually, polar bears are likely to experience increasingly stressful conditions and higher mortality rates. Science , this issue p. 568 ; see also p. 514

Synchrony between reproduction and food availability is important in mammals due to the high energetic costs of gestation and lactation. Female polar bears (Ursus maritimus) must accumulate sufficient energy reserves during spring through autumn to produce and nurse cubs during the winter months in snow dens. Adequate time in a den is important to optimize cub development for withstanding harsh Arctic spring conditions and to synchronize emergence with peak prey availability, which occurs in May and June. During 1985–2013, den phenology was investigated using temperature data collected on satellite collars deployed on adult female polar bears in the southern Beaufort Sea (SB) and Chukchi Sea (CS). We examined relationships between den phenology, reproductive success (cub production and post-emergence survival), and environmental factors (weather and sea-ice conditions). Females observed with cubs emerged later and remained in dens on average 15.0 ± 7.6 (SE) days longer than females seen without cubs. Females occupying land-based dens, where estimated snowfall was greater, had higher reproductive success. Recently, female polar bears have increased land-based denning in the SB. Females in CS emerged later from dens than SB females, consistent with better female body condition and higher cub survival in the CS. During years with a greater area of autumn sea ice, reproductive success was higher at land-based versus sea-ice dens, suggesting continued decline in sea ice could negatively affect recruitment. However, further research is needed to better understand mechanistic relationships. Because females emerging later from dens had higher reproductive success, den duration could be a useful metric in population monitoring.

Plasticity in the physiological and behavioural responses of animals to prolonged food shortages may determine the persistence of species under climate warming. This is particularly applicable for species that can “adaptively fast” by conserving protein to protect organ function while catabolizing endogenous tissues. Some Ursids, including polar bears (Ursus maritimus), adaptively fast during winter hibernation—and it has been suggested that polar bears also employ this strategy during summer. We captured 57 adult female polar bears in the Southern Beaufort Sea (SBS) during summer 2008 and 2009 and measured blood variables that indicate feeding, regular fasting, and adaptive fasting. We also assessed tissue δ13C and δ15N to infer diet, and body condition via mass and length. We found that bears on shore maintained lipid and protein stores by scavenging on bowhead whale (Balaena mysticetus) carcasses from human harvest, while those that followed the retreating sea ice beyond the continental shelf were food deprived. They had low ratios of blood urea to creatinine (U:C), normally associated with adaptive fasting. However, they also exhibited low albumin and glucose (indicative of protein loss) and elevated alanine aminotransferase and ghrelin (which fall during adaptive fasting). Thus, the ~ 70% of the SBS subpopulation that spends summer on the ice experiences more of a regular, rather than adaptive, fast. This fast will lengthen as summer ice declines. The resulting protein loss prior to winter could be a mechanism driving the reported correlation between summer ice and polar bear reproduction and survival in the SBS.

The effects of declining Arctic sea ice on local ecosystem productivity are not well understood but have been shown to vary inter-specifically, spatially, and temporally. Because marine mammals occupy upper trophic levels in Arctic food webs, they may be useful indicators for understanding variation in ecosystem productivity. Polar bears (Ursus maritimus) are apex predators that primarily consume benthic and pelagic-feeding ice-associated seals. As such, their productivity integrates sea ice conditions and the ecosystem supporting them. Declining sea ice availability has been linked to negative population effects for polar bears but does not fully explain observed population changes. We examined relationships between spring foraging success of polar bears and sea ice conditions, prey productivity, and general patterns of ecosystem productivity in the Beaufort and Chukchi Seas (CSs). Fasting status (≥7 days) was estimated using serum urea and creatinine levels of 1,448 samples collected from 1,177 adult and subadult bears across three subpopulations. Fasting increased in the Beaufort Sea between 1983-1999 and 2000-2016 and was related to an index of ringed seal body condition. This change was concurrent with declines in body condition of polar bears and observed changes in the diet, condition and/or reproduction of four other vertebrate consumers within the food chain. In contrast, fasting declined in CS polar bears between periods and was less common than in the two Beaufort Sea subpopulations consistent with studies demonstrating higher primary productivity and maintenance or improved body condition in polar bears, ringed seals, and bearded seals despite recent sea ice loss in this region. Consistency between regional and temporal variation in spring polar bear fasting and food web productivity suggests that polar bears may be a useful indicator species. Furthermore, our results suggest that spatial and temporal ecological variation is important in affecting upper trophic-level productivity in these marine ecosystems.

Recent reductions in thickness and extent have increased drift rates of Arctic sea ice. Increased ice drift could significantly affect the movements and the energy balance of polar bears (Ursus maritimus) which forage, nearly exclusively, on this substrate. We used radio-tracking and ice drift data to quantify the influence of increased drift on bear movements, and we modeled the consequences for energy demands of adult females in the Beaufort and Chukchi seas during two periods with different sea ice characteristics. Westward and northward drift of the sea ice used by polar bears in both regions increased between 1987-1998 and 1999-2013. To remain within their home ranges, polar bears responded to the higher westward ice drift with greater eastward movements, while their movements north in the spring and south in fall were frequently aided by ice motion. To compensate for more rapid westward ice drift in recent years, polar bears covered greater daily distances either by increasing their time spent active (7.6%-9.6%) or by increasing their travel speed (8.5%-8.9%). This increased their calculated annual energy expenditure by 1.8%-3.6% (depending on region and reproductive status), a cost that could be met by capturing an additional 1-3 seals/year. Polar bears selected similar habitats in both periods, indicating that faster drift did not alter habitat preferences. Compounding reduced foraging opportunities that result from habitat loss; changes in ice drift, and associated activity increases, likely exacerbate the physiological stress experienced by polar bears in a warming Arctic.

Greenhouse‐gas‐induced warming in the Arctic has caused declines in sea ice extent and changed its composition, raising concerns by all circumpolar nations for polar bear conservation. Negative impacts have been observed in three well‐studied polar bear subpopulations. Most subpopulations, however, receive little or no direct monitoring, hence, resource selection functions (RSF) may provide a useful proxy of polar bear distributions. However, the efficacy of RSFs constructed from past data, that is, reference RSFs, may be degraded under contemporary conditions, especially in a rapidly changing environment. We assessed published Arctic‐wide reference RSFs using tracking data from adult female polar bears captured in the Beaufort Sea. We compared telemetry‐derived seasonal distributions of polar bears to RSF‐defined optimal sea ice habitat during the period of RSF model development, 1985–1995, and two subsequent periods with diminished sea ice: 1996–2006 and 2007–2016. From these comparisons, we assessed the applicability of the reference RSFs for contemporary polar bear conservation. In the two decades following the 1985–1995 reference period, use and availability of optimal habitat by polar bears declined during the ice melt, ice minimum, and ice growth seasons. During the ice maximum season (i.e., winter), polar bears used the best habitat available, which changed relatively little across the three decades of study. During the ice melt, ice minimum, and ice growth seasons, optimal habitat in areas used by polar bears decreased and was displaced north and east of the Alaska Beaufort Sea coast. As optimal habitat diminished in these seasons, polar bears expanded their range and occupied greater areas of suboptimal habitat. Synthesis and applications: Sea ice declines due to climate change continue to challenge polar bears and their conservation. The distribution of Southern Beaufort Sea polar bears remained similar during the ice maximum season, so the reference RSFs developed from data collected >20 years ago continue to accurately model their winter distribution. In contrast, reference RSFs for the ice transitional and minimum seasons showed diminished predictive efficacy but were useful in revealing that contemporary polar bears have been increasingly forced to use suboptimal habitats during those seasons.

Climate change is altering the distribution of some wildlife species while warming temperatures are facilitating the northward expansion of pathogens, potentially increasing disease risk. Melting of Arctic sea ice is increasingly causing polar bears (Ursus maritimus) of the southern Beaufort Sea (SBS) to spend summer on land, where they may encounter novel pathogens. Here, we tested whether SBS polar bears on shore during summer exhibited greater immune system activity than bears remaining on the sea ice. In addition, we tested whether the type of immune response correlated with body condition, because adaptive responses (slowly developing defenses against specific pathogens) often require less energy than innate responses (rapid defenses not based on pathogen identity). After accounting for body condition, we found that polar bears on shore exhibited higher total white blood cell counts, neutrophils, and monocytes than bears on the ice, suggesting more infections. Lymphocytes, eosinophils, basophils, and globulins did not differ. C-reactive protein, an indicator of inflammation, also did not differ between habitats. Body condition was associated with variables indicative of both innate and adaptive immunity, suggesting that neither response was uniquely limited by energy resources. Our data indicate that as more polar bears spend longer periods of time on shore, they may experience more infections. We encourage continued health monitoring of this species and studies of the long-term fitness consequences from disease.

Not that unusual after all As polar ice recedes, polar bears are facing a changed habitat with reduced summer foraging opportunities. It has been hypothesized that they might be able to resist summer food shortages by reducing their metabolic needs in a sort of “walking hibernation.” Whiteman et al. monitored energy expenditure in polar bears both on and off the ice and found energy reductions, but that these were more akin to normal mammalian fasting levels. Thus, it appears that polar bears have no energetic protections against reduced summer food supplies and will face increasing starvation threats if summer foraging habitats continue to decline. Science , this issue p. 295

In the southern Beaufort Sea of the United States and Canada, prior investigations have linked declines in summer sea ice to reduced physical condition, growth, and survival of polar bears (Ursus maritimus). Combined with projections of population decline due to continued climate warming and the ensuing loss of sea ice habitat, those findings contributed to the 2008 decision to list the species as threatened under the U.S. Endangered Species Act. Here, we used mark-recapture models to investigate the population dynamics of polar bears in the southern Beaufort Sea from 2001 to 2010, years during which the spatial and temporal extent of summer sea ice generally declined. Low survival from 2004 through 2006 led to a 25-50% decline in abundance. We hypothesize that low survival during this period resulted from (1) unfavorable ice conditions that limited access to prey during multiple seasons; and possibly, (2) low prey abundance. For reasons that are not clear, survival of adults and cubs began to improve in 2007 and abundance was comparatively stable from 2008 to 2010, with ~900 bears in 2010 (90% CI 606-1212). However, survival of subadult bears declined throughout the entire period. Reduced spatial and temporal availability of sea ice is expected to increasingly force population dynamics of polar bears as the climate continues to warm. However, in the short term, our findings suggest that factors other than sea ice can influence survival. A refined understanding of the ecological mechanisms underlying polar bear population dynamics is necessary to improve projections of their future status and facilitate development of management strategies.

Polar bears experience skeletal muscle atrophy during winter whether they hibernate or not. Bears spending summer on sea ice in the Arctic basin undergo a second period of atrophy caused by fasting, whereas bears with shore access to adequate, alternative food do not. Summer atrophy may influence fitness., When reducing activity and using stored energy during seasonal food shortages, animals risk degradation of skeletal muscles, although some species avoid or minimize the resulting atrophy while experiencing these conditions during hibernation. Polar bears may be food deprived and relatively inactive during winter (when pregnant females hibernate and hunting success declines for other demographic groups) as well as summer (when sea ice retreats from key foraging habitats). We investigated muscle atrophy in samples of biceps femoris collected from free-ranging polar bears in the Southern Beaufort Sea (SBS) throughout their annual cycle. Atrophy was most pronounced in April–May as a result of food deprivation during the previous winter, with muscles exhibiting reduced protein concentration, increased water content, and lower creatine kinase mRNA. These animals increased feeding and activity in spring (when seal prey becomes more available), initiating a period of muscle recovery. During the following ice melt of late summer, ~30% of SBS bears abandon retreating sea ice for land; in August, these ‘shore’ bears exhibited no muscle atrophy, indicating that they had fully recovered from winter food deprivation. These individuals subsequently scavenged whale carcasses deposited by humans and by October, had retained good muscle condition. In contrast, ~70% of SBS bears follow the ice north in late summer, into deep water with less prey. These ‘ice’ bears fast; by October, they exhibited muscle protein loss and rapid changes in myosin heavy-chain isoforms in response to reduced activity. These findings indicate that, unlike other bears during winter hibernation, polar bears without food in summer cannot mitigate atrophy. Consequently, prolonged summer fasting resulting from climate change-induced ice loss creates a risk of greater muscle atrophy and reduced abilities to travel and hunt.

The polar bear (Ursus maritimus) is an obligate apex predator of Arctic sea ice and as such can be affected by climate warming-induced changes in the extent and composition of pack ice and its impacts on their seal prey. Sea ice declines have negatively impacted some polar bear subpopulations through reduced energy input because of loss of hunting habitats, higher energy costs due to greater ice drift, ice fracturing and open water, and ultimately greater challenges to recruit young. Projections made from the output of global climate models suggest that polar bears in peripheral Arctic and sub-Arctic seas will be reduced in numbers or become extirpated by the end of the twenty-first century if the rate of climate warming continues on its present trajectory. The same projections also suggest that polar bears may persist in the high-latitude Arctic where heavy multiyear sea ice that has been typical in that region is being replaced by thinner annual ice. Underlying physical and biological oceanography provides clues as to why polar bear in some regions are negatively impacted, while bears in other regions have shown no apparent changes. However, continued declines in sea ice will eventually challenge the survival of polar bears and efforts to conserve them in all regions of the Arctic.

Effective conservation planning requires understanding and ranking threats to wildlife populations. We developed a Bayesian network model to evaluate the relative influence of environmental and anthropogenic stressors, and their mitigation, on the persistence of polar bears (Ursus maritimus). Overall sea ice conditions, affected by rising global temperatures, were the most influential determinant of population outcomes. Accordingly, unabated rise in atmospheric greenhouse gas (GHG) concentrations was the dominant influence leading to worsened population outcomes, with polar bears in three of four ecoregions reaching a dominant probability of decreased or greatly decreased by the latter part of this century. Stabilization of atmospheric GHG concentrations by mid‐century delayed the greatly reduced state by ≈25 yr in two ecoregions. Prompt and aggressive mitigation of emissions reduced the probability of any regional population becoming greatly reduced by up to 25%. Marine prey availability, linked closely to sea ice trend, had slightly less influence on outcome state than sea ice availability itself. Reduced mortality from hunting and defense of life and property interactions resulted in modest declines in the probability of a decreased or greatly decreased population outcome. Minimizing other stressors such as trans‐Arctic shipping, oil and gas exploration, and contaminants had a negligible effect on polar bear outcomes, although the model was not well‐informed with respect to the potential influence of these stressors. Adverse consequences of loss of sea ice habitat became more pronounced as the summer ice‐free period lengthened beyond four months, which could occur in most of the Arctic basin after mid‐century if GHG emissions are not promptly reduced. Long‐term conservation of polar bears would be best supported by holding global mean temperature to ≤ 2°C above preindustrial levels. Until further sea ice loss is stopped, management of other stressors may serve to slow the transition of populations to progressively worsened outcomes, and improve the prospects for their long‐term persistence.

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.

Polar bears (Ursus maritimus) occupy remote regions that are characterized by harsh weather and limited access. Polar bear populations can only persist where temporal and spatial availability of sea ice provides adequate access to their marine mammal prey. Observed declines in sea ice availability will continue as long as greenhouse gas concentrations rise. At the same time, human intrusion and pollution levels in the Arctic are expected to increase. A circumpolar understanding of the cumulative impacts of current and future stressors is lacking, long-term trends are known from only a few subpopulations, and there is no globally coordinated effort to monitor effects of stressors. Here, we describe a framework for an integrated circumpolar monitoring plan to detect ongoing patterns, predict future trends, and identify the most vulnerable polar bear subpopulations. We recommend strategies for monitoring subpopulation abundance and trends, reproduction, survival, ecosystem change, human-caused morta...

Polar bears ( Ursus maritimus Phipps, 1774) depend on sea ice for catching marine mammal prey. Recent sea-ice declines have been linked to reductions in body condition, survival, and population size. Reduced foraging opportunity is hypothesized to be the primary cause of sea-ice-linked declines, but the costs of travel through a deteriorated sea-ice environment also may be a factor. We used movement data from 52 adult female polar bears wearing Global Positioning System (GPS) collars, including some with dependent young, to document long-distance swimming (>50 km) by polar bears in the southern Beaufort and Chukchi seas. During 6 years (2004–2009), we identified 50 long-distance swims by 20 bears. Swim duration and distance ranged from 0.7 to 9.7 days (mean = 3.4 days) and 53.7 to 687.1 km (mean = 154.2 km), respectively. Frequency of swimming appeared to increase over the course of the study. We show that adult female polar bears and their cubs are capable of swimming long distances during periods when extensive areas of open water are present. However, long-distance swimming appears to have higher energetic demands than moving over sea ice. Our observations suggest long-distance swimming is a behavioral response to declining summer sea-ice conditions.

Polar bears (Ursus maritimus) prefer to live on Arctic sea ice but may swim between ice floes or between sea ice and land. Although anecdotal observations suggest that polar bears are capable of swimming long distances, no data have been available to describe in detail long distance swimming events or the physiological and reproductive consequences of such behavior. Between an initial capture in late August and a recapture in late October 2008, a radio-collared adult female polar bear in the Beaufort Sea made a continuous swim of 687 km over 9 days and then intermittently swam and walked on the sea ice surface an additional 1,800 km. Measures of movement rate, hourly activity, and subcutaneous and external temperature revealed distinct profiles of swimming and walking. Between captures, this polar bear lost 22% of her body mass and her yearling cub. The extraordinary long distance swimming ability of polar bears, which we confirm here, may help them cope with reduced Arctic sea ice. Our observation, however, indicates that long distance swimming in Arctic waters, and travel over deep water pack ice, may result in high energetic costs and compromise reproductive fitness.

On the basis of projected losses of their essential sea-ice habitats, a United States Geological Survey research team concluded in 2007 that two-thirds of the world's polar bears (Ursus maritimus) could disappear by mid-century if business-as-usual greenhouse gas emissions continue. That projection, however, did not consider the possible benefits of greenhouse gas mitigation. A key question is whether temperature increases lead to proportional losses of sea-ice habitat, or whether sea-ice cover crosses a tipping point and irreversibly collapses when temperature reaches a critical threshold. Such a tipping point would mean future greenhouse gas mitigation would confer no conservation benefits to polar bears. Here we show, using a general circulation model, that substantially more sea-ice habitat would be retained if greenhouse gas rise is mitigated. We also show, with Bayesian network model outcomes, that increased habitat retention under greenhouse gas mitigation means that polar bears could persist throughout the century in greater numbers and more areas than in the business-as-usual case. Our general circulation model outcomes did not reveal thresholds leading to irreversible loss of ice; instead, a linear relationship between global mean surface air temperature and sea-ice habitat substantiated the hypothesis that sea-ice thermodynamics can overcome albedo feedbacks proposed to cause sea-ice tipping points. Our outcomes indicate that rapid summer ice losses in models and observations represent increased volatility of a thinning sea-ice cover, rather than tipping-point behaviour. Mitigation-driven Bayesian network outcomes show that previously predicted declines in polar bear distribution and numbers are not unavoidable. Because polar bears are sentinels of the Arctic marine ecosystem and trends in their sea-ice habitats foreshadow future global changes, mitigating greenhouse gas emissions to improve polar bear status would have conservation benefits throughout and beyond the Arctic.

Projections of polar bear (Ursus maritimus) sea ice habitat distribution in the polar basin during the 21st century were developed to understand the consequences of anticipated sea ice reductions on polar bear populations. We used location data from satellite- collared polar bears and environmental data (e.g., bathymetry, distance to coastlines, and sea ice) collected from 1985 to 1995 to build resource selection functions (RSFs). RSFs described habitats that polar bears preferred in summer, autumn, winter, and spring. When applied to independent data from 1996 to 2006, the RSFs consistently identified habitats most frequently used by polar bears. We applied the RSFs to monthly maps of 21st-century sea ice concentration projected by 10 general circulation models (GCMs) used in the Intergovern- mental Panel of Climate Change Fourth Assessment Report, under the A1B greenhouse gas forcing scenario. Despite variation in their projections, all GCMs indicated habitat losses in the polar basin during the 21st century. Losses in the highest-valued RSF habitat (optimal habitat) were greatest in the southern seas of the polar basin, especially the Chukchi and Barents seas, and least along the Arctic Ocean shores of Banks Island to northern Greenland. Mean loss of optimal polar bear habitat was greatest during summer; from an observed 1.0 million km 2 in 1985-1995 (baseline) to a projected multi-model mean of 0.32 million km 2 in 2090-2099 (� 68% change). Projected winter losses of polar bear habitat were less: from 1.7 million km 2 in 1985-1995 to 1.4 million km 2 in 2090-2099 (� 17% change). Habitat losses based on GCM multi-model means may be conservative; simulated rates of habitat loss during 1985-2006 from many GCMs were less than the actual observed rates of loss. Although a reduction in the total amount of optimal habitat will likely reduce polar bear populations, exact relationships between habitat losses and population demographics remain unknown. Density and energetic effects may become important as polar bears make long-distance annual migrations from traditional winter ranges to remnant high-latitude summer sea ice. These impacts will likely affect specific sex and age groups differently and may ultimately preclude bears from seasonally returning to their traditional ranges.

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.

Although sea ice loss is the primary threat to polar bears (Ursus maritimus), little can be done to mitigate its effects without global efforts to reduce greenhouse gas emissions. Other factors, however, could exacerbate the impacts of sea ice loss on polar bears, such as exposure to increased industrial activity. The Arctic Ocean has enormous oil and gas potential, and its development is expected to increase in the coming decades. Estimates of polar bear resource selection will inform managers how bears use areas slated for oil development and to help guide conservation planning. We estimated temporally-varying resource selection patterns for non-denning adult female polar bears in the Chukchi Sea population (2008–2012) at two scales (i.e., home range and weekly steps) to identify factors predictive of polar bear use throughout the year, before any offshore development. From the best models at each scale, we estimated scale-integrated resource selection functions to predict polar bear space use across th...

Context The potential for research methods to affect wildlife is an increasing concern among both scientists and the public. This topic has a particular urgency for polar bears because additional research is needed to monitor and understand population responses to rapid loss of sea ice habitat. Aims This study used data collected from polar bears sampled in the Alaska portion of the southern Beaufort Sea to investigate the potential for capture to adversely affect behaviour and vital rates. We evaluated the extent to which capture, collaring and handling may influence activity and movement days to weeks post-capture, and body mass, body condition, reproduction and survival over 6 months or more. Methods We compared post-capture activity and movement rates, and relationships between prior capture history and body mass, body condition and reproductive success. We also summarised data on capture-related mortality. Key results Individual-based estimates of activity and movement rates reached near-normal levels within 2–3 days and fully normal levels within 5 days post-capture. Models of activity and movement rates among all bears had poor fit, but suggested potential for prolonged, lower-level rate reductions. Repeated captures was not related to negative effects on body condition, reproduction or cub growth or survival. Capture-related mortality was substantially reduced after 1986, when immobilisation drugs were changed, with only 3 mortalities in 2517 captures from 1987–2013. Conclusions Polar bears in the southern Beaufort Sea exhibited the greatest reductions in activity and movement rates 3.5 days post-capture. These shorter-term, post-capture effects do not appear to have translated into any long-term effects on body condition, reproduction, or cub survival. Additionally, collaring had no effect on polar bear recovery rates, body condition, reproduction or cub survival. Implications This study provides empirical evidence that current capture-based research methods do not have long-term implications, and are not contributing to observed changes in body condition, reproduction or survival in the southern Beaufort Sea. Continued refinement of capture protocols, such as the use of low-impact dart rifles and reversible drug combinations, might improve polar bear response to capture and abate short-term reductions in activity and movement post-capture.