The post-emergence period for denning Polar Bears: phenology and influence on cub survival.

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.

Diet energy density estimated from isotopes in predator hair associated with survival, habitat, and population dynamics.

Sea ice loss is fundamentally altering the Arctic marine environment. Yet there is a paucity of data on the adaptability of food webs to ecosystem change, including predator-prey interactions. Polar bears (Ursus maritimus) are an important subsistence resource for Indigenous people and an apex predator that relies entirely on the under-ice food web to meet its energy needs. In this study, we assessed whether polar bears maintained dietary energy density by prey switching in response to spatiotemporal variation in prey availability. We compared the macronutrient composition of diets inferred from stable carbon and nitrogen isotopes in polar bear guard hair (primarily representing summer/fall diet) during periods when bears had low and high survival (2004-2016), between bears that summered on land versus pack ice, and between bears occupying different regions of the Alaskan and Canadian Beaufort Sea. Polar bears consumed diets with lower energy density during periods of low survival, suggesting that concurrent increased dietary proportions of beluga whales (Delphinapterus leucas) did not offset reduced proportions of ringed seals (Pusa hispida). Diets with the lowest energy density and proportions from ringed seal blubber were consumed by bears in the western Beaufort Sea (Alaska) during a period when polar bear abundance declined. Intake required to meet energy requirements of an average free-ranging adult female polar bear was 2.1 kg/day on diets consumed during years with high survival but rose to 3.0 kg/day when survival was low. Although bears that summered onshore in the Alaskan Beaufort Sea had higher-fat diets than bears that summered on the pack ice, access to the remains of subsistence-harvested bowhead whales (Balaena mysticetus) contributed little to improving diet energy density. Because most bears in this region remain with the sea ice year round, prey switching and consumption of whale carcasses onshore appear insufficient to augment diets when availability of their primary prey, ringed seals, is reduced. Our results show that a strong predator-prey relationship between polar bears and ringed seals continues in the Beaufort Sea. The method of estimating dietary blubber using predator hair, demonstrated here, provides a new metric to monitor predator-prey relationships that affect individual health and population demographics.

The acute physiological response of polar bears to helicopter capture.

Many wildlife species are live captured, sampled, and released; for polar bears (Ursus maritimus) capture often requires chemical immobilization via helicopter darting. Polar bears reduce their activity for approximately 4 days after capture, likely reflecting stress recovery. To better understand this stress, we quantified polar bear activity (via collar-mounted accelerometers) and body temperature (via loggers in the body core [Tabd] and periphery [Tper]) during 2-6 months of natural behavior, and during helicopter recapture and immobilization. Recapture induced bouts of peak activity higher than those that occurred during natural behavior for 2 of 5 bears, greater peak Tper for 3 of 6 bears, and greater peak Tabd for 1 of 6 bears. High body temperature (>39.0°C) occurred in Tper for 3 of 6 individuals during recapture and 6 of 6 individuals during natural behavior, and in Tabd for 2 of 6 individuals during recapture and 3 of 6 individuals during natural behavior. Measurements of Tabd and Tper correlated with rectal temperatures measured after immobilization, supporting the use of rectal temperatures for monitoring bear response to capture. Using a larger dataset (n = 66 captures), modeling of blood biochemistry revealed that maximum ambient temperature during recapture was associated with a stress leukogram (7-26% decline in percent lymphocytes, 12-21% increase in percent neutrophils) and maximum duration of helicopter operations had a similar but smaller effect. We conclude that polar bear activity and body temperature during helicopter capture are similar to that which occurs during the most intense events of natural behavior; high body temperature, especially in warm capture conditions, is a key concern; additional study of stress leukograms in polar bears is needed; and additional data collection regarding capture operations would be useful.

Survival and abundance of polar bears in Alaska's Beaufort Sea, 2001-2016.

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.

Identifying reliable indicators of fitness in polar bears.

Animal structural body size and condition are often measured to evaluate individual health, identify responses to environmental change and food availability, and relate food availability to effects on reproduction and survival. A variety of condition metrics have been developed but relationships between these metrics and vital rates are rarely validated. Identifying an optimal approach to estimate the body condition of polar bears is needed to improve monitoring of their response to decline in sea ice habitat. Therefore, we examined relationships between several commonly used condition indices (CI), body mass, and size with female reproductive success and cub survival among polar bears (Ursus maritimus) measured in two subpopulations over three decades. To improve measurement and application of morphometrics and CIs, we also examined whether CIs are independent of age and structural size-an important assumption for monitoring temporal trends-and factors affecting measurement precision and accuracy. Maternal CIs and mass measured the fall prior to denning were related to cub production. Similarly, maternal CIs, mass, and length were related to the mass of cubs or yearlings that accompanied her. However, maternal body mass, but not CIs, measured in the spring was related to cub production and only maternal mass and length were related to the probability of cub survival. These results suggest that CIs may not be better indicators of fitness than body mass in part because CIs remove variation associated with body size that is important in affecting fitness. Further, CIs exhibited variable relationships with age for growing bears and were lower for longer bears despite body length being related to cub survival and female reproductive success. These results are consistent with findings from other species indicating that body mass is a useful metric to link environmental conditions and population dynamics.

The seasonal energetic landscape of an apex marine carnivore, the polar bear.

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.

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

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.

Accounting for phenology in the analysis of animal movement.

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.

Heightened Immune System Function in Polar Bears Using Terrestrial Habitats.

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.

Phenotypic plasticity and climate change: can polar bears respond to longer Arctic summers with an adaptive fast?

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.

Spring fasting behavior in a marine apex predator provides an index of ecosystem productivity.

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.

Polar bears experience skeletal muscle atrophy in response to food deprivation and reduced activity in winter and summer.

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.

Increased Arctic sea ice drift alters adult female polar bear movements and energetics.

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.

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.

Polar bear population dynamics in the southern Beaufort Sea during a period of sea ice decline.

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.

Greenhouse gas mitigation can reduce sea-ice loss and increase polar bear persistence.

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.