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1. Quantifying the diet composition of apex marine predators such as killer whales (Orcinus orca) is critical to assessing their food web impacts. Yet, with few exceptions, the feeding ecology of these apex predators remains poorly understood. 2. Here, we use our newly validated quantitative fatty acid signature analysis (QFASA) approach on nearly 200 killer whales and over 900 potential prey to model their diets across the 5000 km span of the North Atlantic. 3. Diet estimates show that killer whales mainly consume other whales in the western North Atlantic (Canadian Arctic, Eastern Canada), seals in the mid-North Atlantic (Greenland), and fish in the eastern North Atlantic (Iceland, Faroe Islands, Norway). Nonetheless, diet estimates also varied widely among individuals within most regions. This level of inter-individual feeding variation should be considered for future ecological studies focusing on killer whales in the North Atlantic and other oceans. 4. These estimates reveal remarkable population- and individual-level variation in the trophic ecology of these killer whales, which can help to assess how their predation impacts community and ecosystem dynamics in changing North Atlantic marine ecosystems. 5. This new approach provides researchers with an invaluable tool to study the feeding ecology of oceanic top predators.

The monitoring of legacy persistent organic pollutants (POPs) in blubber of key sentinel marine mammal species has been conducted using established techniques for decades. Although these methods for polychlorinated biphenyl (PCB) and organochlorine (OC) pesticide determination provide accurate and reproducible results, they possess some drawbacks in terms of cost, time, and a need for large volumes of toxic solvents. QuEChERS (quick, easy, cheap, effective, rugged, and safe) extractions may help address these issues, but have not been applied to marine mammal blubber/adipose. As such, our aim was to develop, validate, and apply a QuEChERS method for the extraction of PCB and OC contaminants in marine mammal blubber. First, we tested multiple solid-phase extraction and clean-up steps to find the approach that provided the cleanest extracts along with consistent and acceptable analyte recovery, accuracy, and precision. QuEChERS extractions followed by two enhanced matrix removal-lipid (EMR-lipid), one primary-secondary amine (PSA), and one silica gel clean-up showed the highest matrix removal and acceptable recoveries of spiked internal (62–97%) and external standards (61–94%). Solvent usage was reduced by ∼393% and extraction time was reduced by ∼25% (from 16 to 12 h). Next, the method was validated using standard reference material (SRM) NIST 1945. Recovery experiments on SRM (n = 5) showed acceptable recovery for 76% and 77% of PCBs and OC pesticides, respectively, and high precision for 73% and 69% of PCBs and OCs, respectively. Finally, the method was used on a set of southeast Greenland killer whales (n = 13), with previously published PCB and OC data. Bland-Altman plots indicated good agreement between QuEChERS and current-use methods for ΣPCBs and some OCs with no significant constant or proportional bias. These results demonstrate that this QuEChERS extraction method represents an effective, lower cost alternative to current-use extractions for PCBs and OCs in blubber, and likely other high-lipid samples.

Aim: Identify hotspots and areas of high species richness for Arctic marine mammals. Location: Circumpolar Arctic. Methods: A total of 2115 biologging devices were deployed on marine mammals from 13 species in the Arctic from 2005 to 2019. Getis-Ord Gi* hotspots were calculated based on the number of individuals in grid cells for each species and for phyloge-netic groups (nine pinnipeds, three cetaceans, all species) and areas with high spe-cies richness were identified for summer (Jun-Nov), winter (Dec-May) and the entire year. Seasonal habitat differences among species’ hotspots were investigated using Principal Component Analysis. Results: Hotspots and areas with high species richness occurred within the Arctic continental-shelf seas and within the marginal ice zone, particularly in the “Arctic gateways” of the north Atlantic and Pacific oceans. Summer hotspots were generally found further north than winter hotspots, but there were exceptions to this pattern, including bowhead whales in the Greenland-Barents Seas and species with coastal distributions in Svalbard, Norway and East Greenland. Areas with high species rich-ness generally overlapped high-density hotspots. Large regional and seasonal dif-ferences in habitat features of hotspots were found among species but also within species from different regions. Gap analysis (discrepancy between hotspots and IUCN ranges) identified species and regions where more research is required. Main conclusions: This study identified important areas (and habitat types) for Arctic marine mammals using available biotelemetry data. The results herein serve as a benchmark to measure future distributional shifts. Expanded monitoring and teleme-try studies are needed on Arctic species to understand the impacts of climate change and concomitant ecosystem changes (synergistic effects of multiple stressors). While efforts should be made to fill knowledge gaps, including regional gaps and more com-plete sex and age coverage, hotspots identified herein can inform management ef-forts to mitigate the impacts of human activities and ecological changes, including creation of protected areas.

Two major oceanographic changes have recently propagated through several trophic levels in coastal areas of Southeast Greenland (SEG). Firstly, the amount of drift-ice exported from the Fram Strait and transported with the East Greenland Current (EGC) has decreased significantly over the past two decades, and a main tipping element (summer sea ice) has virtually disappeared since 2003 leading to a regime shift in oceanographic and ecological conditions in the region. The following 20-year period with low or no coastal sea ice is unique in the 200-year history of ice observations in the region, and the regime shift is also obvious in the volume of ice export through the Fram Strait after 2013. In the same period, the temperature of the EGC south of 73.5 N has increased significantly (>2°C) since 1980. Secondly, the warm Irminger Current, which advects warm, saline Atlantic Water into the region, has become warmer since 1990. The lack of pack ice in summer together with a warming ocean generatedcascading effects on the ecosystem in SEG that are manifested in a changed fishfauna with an influx of boreal species in the south and the subarctic capelin further north. At higher trophic levels there has been an increase in the abundance of severalboreal cetaceans (humpback, fin, killer, and pilot whales and dolphins) that are eithernew to this area or occur in historically large numbers. It is estimated that the new cetacean species in SEG are responsible for an annual predation level of 700,000 tonsof fish. In addition, predation on krill species is estimated at >1,500,000 tons mainlyconsumed by fin whales. Simultaneously, there has been a reduction in the abundance and catches of narwhals and walruses in SEG and it is suggested that these specieshave been impacted by the habitat changes.

There has been a considerable number of reports on Hg concentrations in Arctic mammals since the last Arctic Monitoring and Assessment Programme (AMAP) effort to review biological effects of the exposure to mercury (Hg) in Arctic biota in 2010 and 2018. Here, we provide an update on the state of the knowledge of health risk associated with Hg concentrations in Arctic marine and terrestrial mammal species. Using available population-specific data post-2000, our ultimate goal is to provide an updated evidence-based estimate of the risk for adverse health effects from Hg exposure in Arctic mammal species at the individual and population level. Tissue residues of Hg in 13 species across the Arctic were classified into five risk categories (from No risk to Severe risk) based on critical tissue concentrations derived from experimental studies on harp seals and mink. Exposure to Hg lead to low or no risk for health effects in most populations of marine and terrestrial mammals, however, subpopulations of polar bears, pilot whales, narwhals, beluga and hooded seals are highly exposed in geographic hotspots raising concern for Hg-induced toxicological effects. About 6% of a total of 3500 individuals, across different marine mammal species, age groups and regions, are at high or severe risk of health effects from Hg exposure. The corresponding figure for the 12 terrestrial species, regions and age groups was as low as 0.3% of a total of 731 individuals analyzed for their Hg loads. Temporal analyses indicated that the proportion of polar bears at low or moderate risk has increased in East/West Greenland and Western Hudson Bay, respectively. However, there remain numerous knowledge gaps to improve risk assessments of Hg exposure in Arctic mammalian species, including the establishment of improved concentration thresholds and upscaling to the assessment of population-level effects.

The study was partly conducted under a Villum Foundation Young Investigator grant (19114) awarded to IM, which also supports XL, as well as the BONUS BALTHEALTH project (Art. 185) awarded to RDI, AG, TH and MTO. BONUS BALTHEALTH was funded jointly by the EU, Innovation Fund Denmark (grants 6180-00001B and 6180-00002B), Forschungszentrum Jülich GmbH, German Federal Ministry of Education and Research (grant FKZ 03F0767A), Academy of Finland (grant 311966) and Swedish Foundation for Strategic Environmental Research (MISTRA). The harbour seal (Phoca vitulina) is the most widely distributed pinniped, occupying a wide variety of habitats and climatic zones across the Northern Hemisphere. Intriguingly, the harbour seal is also one of the most philopatric seals, raising questions as to how it colonised virtually the whole of the Northern Hemisphere. To shed light on the origin, remarkable range expansion, population structure and genetic diversity of this species, we used genotyping-by-sequencing to analyse ~13,500 biallelic SNPs from 286 individuals sampled from 22 localities across the species' range. Our results point to a Northeast Pacific origin, colonisation of the North Atlantic via the Canadian Arctic, and subsequent stepping-stone range expansions across the North Atlantic from North America to Europe, accompanied by a successive loss of genetic diversity. Our analyses further revealed a deep divergence between modern North Pacific and North Atlantic harbour seals, with finer-scale genetic structure at regional and local scales consistent with strong philopatry. The study provides new insights into the harbour seal's remarkable ability to colonise and adapt to a wide range of habitats. Furthermore, it has implications for current harbour seal subspecies delineations and highlights the need for international and national red lists and management plans to ensure the protection of genetically and demographically isolated populations. Publisher PDF

The ringed seal is a small phocid seal that has a northern circumpolar distribu-tion. It has long been recognized that body size is variable in ringed seals, and it has been suggested that ecotypes that differ in size exist. This study explores patterns of body size (length and girth) and age-at-maturity across most of the Arctic subspecies’ range using morphometric data from 35 sites. Asymptotic lengths varied from 113 to 151 cm, with sites falling into five distinct size clus-ters (for each sex). Age-at-maturity ranged from 3.1 to 7.4 years, with sites that had early ages of sexual maturity generally having small length-at-maturity and small final body length. The sexes differed in length at some sites, but not in a consistent pattern of dimorphism. The largest ringed seals occurred in western Greenland and eastern Canada, and the smallest occurred in Alaska and the White Sea. Latitudinal trends occurred only within sites in the eastern Canadian Arctic. Girth (with length and season accounted for) was also highly variable but showed no notable spatial pattern; males tended to be more rotund than females. Genetic studies are needed, starting with the “giants” at Kangia (Greenland) and in northern Canada to determine whether they are genetically distinct ecotypes. Additional research is also needed to understand the ecological linkages that drive the significant regional size differences in ringed seals that were confirmed in this study, and also to understand their implications with respect to potential adaptation to climate change.

Using social media, the Greenland Institute of Natural Resources collected data on the occurrence of pink salmon (Oncorhynchus gorbuscha) in 2019. Eighty‐four pink salmon were reported from 22 locations across Greenland. This comprised 76 specimens from 2019 and 8 specimens from 2013 to 2018. Of these, 12 were caught in fresh water, and a single pink salmon was from the bottom of the Nuuk Fjord near the Kapisillit River – the only known river in Greenland where the Atlantic salmon (Salmo salar) spawn. It is unknown if pink salmon have reproduced in Greenland waters.

Environmental change and increasing levels of human activity are threats to marine mammals in the Arctic. Identifying marine mammal hotspots and areas of high species richness are essential to help guide management and conservation efforts. Herein, space use based on biotelemetric tracking devices deployed on 13 species (ringed seal Pusa hispida, bearded seal Erignathus barbatus, harbour seal Phoca vitulina, walrus Odobenus rosmarus, harp seal Pagophilus groenlandicus, hooded seal Cystophora cristata, polar bear Ursus maritimus, bowhead whale Balaena mysticetus, narwhal Monodon monoceros, white whale Delphinapterus leucas, blue whale Balaenoptera musculus, fin whale Balaenoptera physalus and humpback whale Megaptera novaeangliae; total = 585 individuals) in the Greenland and northern Barents Seas between 2005 and 2018 is reported. Getis-Ord Gi* hotspots were calculated for each species as well as all species combined, and areas of high species richness were identified for summer/autumn (Jun-Dec), winter/spring (Jan-May) and the entire year. The marginal ice zone (MIZ) of the Greenland Sea and northern Barents Sea, the waters surrounding the Svalbard Archipelago and a few Northeast Greenland coastal sites were identified as key marine mammal hotspots and areas of high species richness in this region. Individual hotspots identified areas important for most of the tagged animals, such as common resting, nursing, moulting and foraging areas. Location hotspots identified areas heavily used by segments of the tagged populations, including denning areas for polar bears and foraging areas. The hotspots identified herein are also important habitats for seabirds and fishes, and thus conservation and management measures targeting these regions would benefit multiple groups of Arctic animals.

To support sustainable management of apex predator populations, it is important to estimate population size and understand the drivers of population trends to anticipate the consequences of human decisions. Robust population models are needed, which must be based on realistic biological principles and validated with the best available data. A team of international experts reviewed age-structured models of North Atlantic pinniped populations, including Grey seal (Halichoerus grypus), Harp seal (Pagophilus groenlandicus), and Hooded seal (Cystophora cristata). Statistical methods used to fit such models to data were compared and contrasted. Differences in biological assumptions and model equations were driven by the data available from separate studies, including observation methodology and pre-processing. Counts of pups during the breeding season were used in all models, with additional counts of adults and juveniles available in some. The regularity and frequency of data collection, including survey counts and vital rate estimates, varied. Important differences between the models concerned the nature and causes of variation in vital rates (age-dependent survival and fecundity). Parameterisation of age at maturity was detailed and time-dependent in some models and simplified in others. Methods for estimation of model parameters were reviewed and compared. They included Bayesian and maximum likelihood (ML) approaches, implemented via bespoke coding in C, C++, TMB or JAGS. Comparative model runs suggested that as expected, ML-based implementations were rapid and computationally efficient, while Bayesian approaches, which used MCMC or sequential importance sampling, required longer for inference. For grey seal populations in the Netherlands, where preliminary ML-based TMB results were compared with the outputs of a Bayesian JAGS implementation, some differences in parameter estimates were apparent. For these seal populations, further investigations are recommended to explore differences that might result from the modelling framework and model-fitting methodology, and their importance for inference and management advice. The group recommended building on the success of this workshop via continued collaboration with ICES and NAMMCO assessment groups, as well as other experts in the marine mammal modelling community. Specifically, for Northeast Atlantic harp and hooded seal populations, the workshop represents the initial step towards a full ICES benchmark process aimed at revising and evaluating new assessment models. Publisher PDF

Harbor seals (Phoca vitulina) were formerly widely distributed in Greenland, but in most areas, they have now become rare or extinct. In this study, we deployed 15 satellite-linked data-recorders on 12 individual harbor seals captured near the southern tip of Greenland. The tagging site, a small archipelago, turned out to be their main haul-out and molting location. The seals were tagged around the first week of September across 2 years. Six adult seals transmitted during the breeding season (June–July), during which two males and a female stayed near the tagging site, whereas two females and a male made a targeted swim about 250 km northward along the east coast, just prior to parturition and they returned right after the breeding period. Based on behavior (strong site fidelity and abrupt changes in haul-out and diving activities), we determined that all three females gave birth and that parturition happened during 14–21 June. Prolonged haul-out, relating to the molt, started in early August and peaked during 15 August–8 September with an average of 10–11 h haul-out per day. The seals thereafter gradually transitioned toward a winter mode with more deep dives and on average hauling out 4 h every other day. Dives deeper than 400 m made up less than 1% of their dives, but four seals dove to depths in excess of 575 m, with a record depth of 631 m and the longest dives lasting between 20 and 25 min. To our knowledge, these dives are the deepest recorded for harbor seals.

This article describes the relationship between sea ice and marine mammals around Greenland, the possible effects of climate change and the importance of subsistence hunting in modern Greenland. We also summarize the biology, conservation status and utilization of the polar bear, walrus, seal and whale species of Greenland.

Accepted manuscript version. Published version available at https://doi.org/10.1016/j.vetimm.2018.02.005. Licensed CC BY-NC-ND 4.0. Zoonotic infections transmitted from marine mammals to humans in the Baltic and European Arctic are of unknown significance, despite given considerable potential for transmission due to local hunt. Here we present results of an initial screening for Brucella spp. in Arctic and Baltic seal species. Baltic ringed seals (Pusa hispida, n = 12) sampled in October 2015 and Greenland Sea harp seals (Pagophilus groenlandicus, n = 6) and hooded seals (Cystophora cristata, n = 3) sampled in March 2015 were serologically analysed for antibodies against Brucella spp. The serological analyses were performed using the Rose Bengal Test (RBT) followed by a confirmatory testing of RBT-positive samples by a competitive-enzyme linked immunosorbent assay (C-ELISA). Two of the Baltic ringed seals (a juvenile male and a juvenile female) were seropositive thus indicating previous exposure to a Brucella spp. The findings indicate that ringed seals in the Baltic ecosystem may be exposed to and possibly infected by Brucella spp. No seropositive individuals were detected among the Greenland harp and hooded seals. Although our initial screening shows a zoonotic hazard to Baltic locals, a more in-depth epidemiological investigation is needed in order to determine the human risk associated with this.

Environmental variation influences resource distribution, thereby affecting animal movement and foraging decisions. Climate change is altering environmental processes worldwide, but particularly in the Arctic, where changes in the phenology of sea ice have been redistributing resources across space and time. How polar marine predators such as ringed seals Pusa hispida hispida, whose ecology is tightly tied to sea ice, respond to different sea ice dynamics across large spatial scales is generally unknown. Here, behavioural states (resident and traveling) were estimated using state-space models on adult (n = 45) and subadult (n = 85) ringed seal satellite telemetry tracks from 6 Arctic locations. Tagged ringed seals spanned a wide latitudinal (56.54° to 75.58°N) and sea ice phenological range from short (1 to 2 mo) to longer (6 mo) ice-free periods. We assessed the influences of age class and several intra- and inter-annual environmental variables on ringed seal movement ecology. Both adults and subadults spent most of the ice-free season in a resident state (93 and 77%, respectively). A latitudinal gradient was characterised, where longer ice-free seasons and less inter-annual variability in sea ice phenology at lower latitudes were related to ringed seals spending more time in a resident state than their conspecifics at higher latitudes (90 versus 58%, respectively), where the ice-free season was shorter and sea ice phenology between years was less synchronous. Ringed seals are responding to latitudinal differences in sea ice phenology which affect prey distribution, suggesting plasticity in their foraging decisions and spatiotemporal differences in prey distribution across the rapidly changing Arctic.

Determining the male and female representation in zooarchaeological material from hunted animal species is essential, to fully investigate the effects and means of prehistoric hunting practices, and may further provide valuable biological information on past animal life-history, behaviour and demography. However, the fragmented nature of the zooarchaeological record and a lack of clear diagnostic skeletal markers, often prevents such inference. Here, we test the usability of the dog nuclear genome (CanFam3.1) for sex identification of pinnipeds. First, a contemporary sample set (n=72) of ringed seals (Pusa hispida), with known sex was used to test the genetic sex identification method. By quantifying the proportion of X chromosome reads, as the chrX/chr1 ratio, the ratios clustered in two clearly distinguishable sex groups. Of the 72 individuals, 69 were identified to the accurate sex, which proves a high reliability of the genetic method. Second, random down sampling of a subset of the ringed seal samples to different read number, suggests at least 5000 DNA sequence reads mapped to the reference genome as the lower limit for which this method is applicable. Finally, applying this standard, sex identification was successfully carried out on a broad set of ancient pinniped samples, including walruses (Odobenus rosmarus), grey seals (Halichoerus grypus) and harp seals (Pagophilus groenlandicus), which all showed clearly distinct male and female chrX/chr1 ratio groups.

There is increasing evidence that the ∼20 routinely monitored perfluoroalkyl and polyfluoroalkyl substances (PFASs) account for only a fraction of extractable organofluorine (EOF) occurring in the environment. To assess whether PFAS exposure is being underestimated in marine mammals from the Northern Hemisphere, we performed a fluorine mass balance on liver tissues from 11 different species using a combination of targeted PFAS analysis, EOF and total fluorine determination, and suspect screening. Samples were obtained from the east coast United States (US), west and east coast of Greenland, Iceland, and Sweden from 2000 to 2017. Of the 36 target PFASs, perfluorooctane sulfonate (PFOS) dominated in all but one Icelandic and three US samples, where the 7:3 fluorotelomer carboxylic acid (7:3 FTCA) was prevalent. This is the first report of 7:3 FTCA in polar bears (∼1000 ng/g, ww) and cetaceans (

Aim: Climate change is altering marine ecosystems worldwide and is most pronounced in the Arctic. Economic development is increasing leading to more disturbances and pressures on Arctic wildlife. Identifying areas that support higher levels of predator abundance and biodiversity is important for the implementation of targeted conservation measures across the Arctic. Location: Primarily Canadian Arctic marine waters but also parts of the United States, Greenland and Russia. Methods: We compiled the largest data setof existing telemetry data for marine predators in the North American Arctic consisting of 1,283 individuals from 21 species. Data were arranged into four species groups: (a) cetaceans and pinnipeds, (b) polar bears Ursus maritimus (c) seabirds, and (d) fishes to address the following objectives: (a) to identify abundance hotspots for each species group in the summer–autumn and winter–spring; (b) to identify species diversity hotspots across all species groups and extent of overlap with exclusive economic zones; and (c) to perform a gap analysis that assesses amount of overlap between species diversity hotspots with existing protected areas. Results: Abundance and species diversity hotpots during summer–autumn and winter–spring were identified in Baffin Bay, Davis Strait, Hudson Bay, Hudson Strait, Amundsen Gulf, and the Beaufort, Chukchi and Bering seas both within and across species groups. Abundance and species diversity hotpots occurred within the conti -nental slope in summer–autumn and offshore in areas of moving pack ice in winter–spring. Gap analysis revealed that the current level of conservation protection that overlaps species diversity hotspots is low covering only 5% (77,498 km2) in summer–autumn and 7% (83,202 km2) in winter–spring. Main conclusions: We identified several areas of potential importance for Arctic marine predators that could provide policymakers with a starting point for conservation measures given the multitude of threats facing the Arctic. These results are relevant to multilevel and multinational governance to protect this vulnerable ecosystem in our rapidly changing world.

Since the last Arctic Monitoring and Assessment Programme (AMAP) effort to review biological effects of the exposure to organohalogen compounds (OHCs) in Arctic biota, there has been a considerable number of new Arctic effect studies. Here, we provide an update on the state of the knowledge of OHC, and also include mercury, exposure and/or associated effects in key Arctic marine and terrestrial mammal and bird species as well as in fish by reviewing the literature published since the last AMAP assessment in 2010. We aimed at updating the knowledge of how single but also combined health effects are or can be associated to the exposure to single compounds or mixtures of OHCs. We also focussed on assessing both potential individual as well as population health impacts using population-specific exposure data post 2000. We have identified quantifiable effects on vitamin metabolism, immune functioning, thyroid and steroid hormone balances, oxidative stress, tissue pathology, and reproduction. As with the previous assessment, a wealth of documentation is available for biological effects in marine mammals and seabirds, and sentinel species such as the sledge dog and Arctic fox, but information for terrestrial vertebrates and fish remain scarce. While hormones and vitamins are thoroughly studied, oxidative stress, immunotoxic and reproductive effects need further investigation. Depending on the species and population, some OHCs and mercury tissue contaminant burdens post 2000 were observed to be high enough to exceed putative risk threshold levels that have been previously estimated for non-target species or populations outside the Arctic. In this assessment, we made use of risk quotient calculations to summarize the cumulative effects of different OHC classes and mercury for which critical body burdens can be estimated for wildlife across the Arctic. As our ultimate goal is to better predict or estimate the effects of OHCs and mercury in Arctic wildlife at the individual, population and ecosystem level, there remain numerous knowledge gaps on the biological effects of exposure in Arctic biota. These knowledge gaps include the establishment of concentration thresholds for individual compounds as well as for realistic cocktail mixtures that in fact indicate biologically relevant, and not statistically determined, health effects for specific species and subpopulations. Finally, we provide future perspectives on understanding Arctic wildlife health using new in vivo, in vitro, and in silico techniques, and provide case studies on multiple stressors to show that future assessments would benefit from significant efforts to integrate human health, wildlife ecology and retrospective and forecasting aspects into assessing the biological effects of OHC and mercury exposure in Arctic wildlife and fish. © 2019 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).

The importance of the North Water polynya in Smith Sound as an overwintering area for marine mammals has been questioned. One way to address the issue is to assess the abundance of selected marine mammals that are present during winter in the North Water. Visual aerial surveys involving double observer platforms were conducted over the eastern part of the North Water polynya in April 2014. Four species of marine mammals were included in strip-census estimation of abundance. Perception bias was addressed using a double-platform survey protocol, a Chapman mark–recapture estimator for whales, seals and walruses (Odobenus rosmarus) on ice and a mark–recapture distance sampling estimation technique for walruses in water. Availability bias was addressed by correcting abundance estimates by the percentage of time animals detected in water that were available for detection at the surface. The resulting estimates suggested that 2544 walruses (95 % CI 1513–4279), 6005 bearded seals (Erignathus barbatus, 95 % CI 4070–8858), 2324 belugas (Delphinapterus leucas, 95 % CI 968–5575) and 3059 narwhals (Monodon monoceros, 95 % CI 1760–5316) wintered in the eastern part of the North Water polynya in April 2014. The walrus estimate is larger than previous summer estimates, and it emphasizes the importance of the habitat along the Greenland coast as a walrus wintering ground. The estimate of belugas is likely negatively biased due to the partial coverage of the potential habitat. The estimate of narwhals is large compared to the few previous observations of narwhals in winter in the North Water, and it demonstrates that large numbers of narwhals winter there. The overall conclusion is that the North Water is indeed an important wintering area for at least walruses, belugas, narwhals and bearded seals.

This research was supported by grants to J.-P.D. from the Canadian National Science and Engineering Research Council (NSERC) (PGSD3-443700-2013) and Aarhus University’s Graduate School and Science and Technology (GSST) and Department of Bioscience; and by funding from the Danish DANCEA program (MST-112-00171 and MST-112-00199); the Defra, Scottish and Welsh Governments (for CSIP/SMASS/CEFAS); and the Icelandic Research Fund (i. Rannsóknasjóður; grant no. 120248042). B.M. was supported by funding from NERC (grant no. SMRU 10001). This paper is a contribution from the BONUS BALTHEALTH project, which has received funding from BONUS (Art. 185), funded jointly by the EU, Innovation Fund Denmark, Forschungszentrum Jülich GmbH, German Federal Ministry of Education and Research (grant no. FKZ 03F0767A), Academy of Finland (decision no. 311966), and Swedish Foundation for Strategic Environmental Research. Killer whales (Orcinus orca) are among the most highly polychlorinated biphenyl (PCB)–contaminated mammals in the world, raising concern about the health consequences of current PCB exposures. Using an individual-based model framework and globally available data on PCB concentrations in killer whale tissues, we show that PCB-mediated effects on reproduction and immune function threaten the long-term viability of >50% of the world’s killer whale populations. PCB-mediated effects over the coming 100 years predicted that killer whale populations near industrialized regions, and those feeding at high trophic levels regardless of location, are at high risk of population collapse. Despite a near-global ban of PCBs more than 30 years ago, the world’s killer whales illustrate the troubling persistence of this chemical class. Postprint

An increasing number of North Atlantic killer whales Orcinus orca have recently been observed in sub-Arctic and Arctic seas. Within these regions, marine mammal consumption appears frequent relative to the more fish-based diet within traditional habitats. To provide insight into feeding habits of Northward range-expanding killer whales, full-depth blubber fatty acid signatures from 21 free-ranging killer whales from Southeast Greenland, Faroe Islands, and Denmark were compared to those of 4 managed-care killer whales, which were fed a constant, long-term fish diet. We analyzed the entire blubber layer in 10 equal-length subsections to evaluate how fatty acid stratification throughout blubber depth May influence fatty acid-based feeding evaluations. Specific fatty acid markers previously linked to marine mammal feeding in other killer whale populations were significantly higher in free-ranging killer whales relative to managed-care individuals, suggesting that marine mammals represent a time-integrated component of free-ranging killer whales’ diet. Unlike the managed-care whales, fatty acid signatures were highly variable among the free-ranging killer whales, suggesting that either they are generalists or exhibit inter-individual feeding variation. All samples, regardless of origin, showed stratification of fatty acid signatures through blubber layers. Dietary fatty acids generally occurred in higher proportions and were more variable in the innermost layers for the free-ranging whales. These data suggest that superficial blubber sampling through biopsy darting May not capture fine-scale and/or short-term variation in diet, and therefore the sampling approach should be carefully considered in research using fatty acids to evaluate feeding ecology of killer whales and other cetaceans.

Zoonotic infections transmitted from terrestrial and marine mammals to humans in European Arctic are of unknown significance, despite considerable potential for transmission due to local hunt and a rapidly changing environment. As an example, infection with Brucella bacteria may have significant impact on human health due to consumption of raw meat or otherwise contact with tissues and fluids of infected game species such as muskoxen and polar bears. Here, we present serological results for Baffin Bay polar bears (Ursus maritimus) (n = 96) and North East Greenland muskoxen (Ovibos moschatus) (n = 32) for antibodies against Brucella spp. The analysis was a two-step trial initially using the Rose Bengal Test (RBT), followed by confirmative competitive enzyme-linked immunosorbent assays of RBT-positive samples. No muskoxen had antibodies against Brucella spp., while antibodies were detected in six polar bears (6.25%) rendering a seroprevalence in line with previous findings in other Arctic regions. Seropositivity was not related to sex, age or biometrics i.e. size and body condition. Whether Brucella spp. antibodies found in polar bears were due to either prey spill over or true recurrent Brucella spp. infections is unknown. Our results therefore highlight the importance of further research into the zoonotic aspects of Brucella spp. infections, and the impact on wildlife and human health in the Arctic region. Prevalence of antibodies against Brucella spp. in West Greenland polar bears (Ursus maritimus) and East Greenland muskoxen (Ovibos moschatus) © Springer-Verlag GmbH Germany, part of Springer Nature 2018. This is a post-peer-review, pre-copyedit version of an article published in Polar Biology. Locked until 19 March 2019 due to copyright restrictions. The final authenticated version is available online at: https://doi.org/10.1007/s00300-018-2307-4.

Jakobshavn Glacier, west Greenland, has responded to temperature changes in Ilulissat Icefjord, into which it terminates. This study collected hydrographic observations inside Ilulissat Icefjord and from adjacent Disko Bay between 2001 and 2014. The warmest deep Disko Bay waters were blocked by the entrance sill and did not reach Jakobshavn Glacier. In the fjord basin, the summer mean temperature was 2.8°C from 2009 to 2013, excluding 2010, when it was 1°C cooler. Despite this variability, summer potential densities in the basin were in the narrow range of 27.20 ≤ σθ ≤ 27.31 kg m−3, and basin water properties matched those of Disko Bay in this layer each summer. This relation has likely held since at least 1980. Basin waters from 2009 and 2011–13 were therefore similar to those in 1998/99, when Jakobshavn Glacier began to retreat, while basin waters in 2010 were as cool as in the 1980s. The 2010 basin temperature anomaly was advected into Disko Bay, not produced by local atmospheric variability.This anomaly also shows that Ilulissat Icefjord basin waters were renewed annually or faster. Time series fragments inside the fjord did not capture the 2010 anomaly but show that the basin temperatures varied little subannually, outside of summer. Fjord velocity profiles from summer 2013 implied a basin renewal time scale of about 1 month. In model simulations of the fjord circulation, subglacial discharge from Jakobshavn Glacier could drive renewal of the fjord basin over a single summer, while baroclinic forcing from outside the fjord could not, because of the sill at the mouth.

Sightings of killerwhales (Orcinus orca) in Greenland have increased in recent years, coincidentwith sea ice loss. These killerwhales are likely from fish-feeding North Atlantic populations, but may have access tomarinemammal prey in Greenlandic waters, which could lead to increased exposures to biomagnifying contaminants. Most studies on polychlorinated biphenyl (Sigma PCB) and organochlorine (OC) contaminants in killerwhales have used biopsies which may not be representative of contaminant concentrations through the entire blubber depth. Here, we measured PCB and OC concentrations in 10 equal-length blubber sections of 18 killer whales harvested in southeast Greenland (2012-2014), and 3 stranded in the Faroe Islands (2008) and Denmark (2005). Overall, very high concentrations of Sigma PCB, Schlordanes (Sigma CHL), and Sdichlorodiphenyltrichloroethane (Sigma DDT) were found in the southeast Greenland and Denmark individuals (means of similar to 40 to 70 mg kg(-1) lipid weight). These concentrations were higher than in the Faroe Island individuals (means of similar to 2 to 5 mg kg(-1) lipid weight) and above those previously reported for other fish-feeding killerwhales in the North Atlantic, likely in part due to additional feeding onmarinemammals. On a wetweight basis, concentrations of all contaminants were significantly lower in the outermost blubber layer (0.15-0.65 cm) compared to all other layers (p

We quantified blubber concentrations of vitamins A (retinol) and E (alpha-tocopherol) and evaluated associations with persistent organic pollutants (Sigma POPs) in 14 highly-contaminated killer whales (Orcinus orca) sampled in Greenland from 2012 to 2014. We considered the influence of blubber depth, sex/age class and diet (based on biomass % of major fatty acids) in these relationships. Blubber concentrations of vitamin A averaged 34.1 +/- 4.7 mu g g(-1) wet weight (ww) and vitamin E averaged 35.6 +/- 4.4 mu g g(-1) ww. Although overall vitamin A concentrations did not vary between inner (closer to the muscle) and outer (closer to the skin) blubber layer or between sub-adults and adult females, concentrations in the outer layer of sub-adults were lower compared to the outer layer of adult females (p = 0.03). Outer layer may therefore reflect age accumulation of vitamin A, while in the more active inner layer, age effects might be masked by metabolic needs such as lactation. Neither diet nor Sigma POPs affected vitamin A variation, suggesting this vitamin is highly regulated in the body. Given the high exposures in these killer whales, vitamin A might not be a sensitive biomarker for POPs adverse effects. Vitamin E concentrations were significantly higher in inner compared to outer layer (p

[1] Interaction of warm, Atlantic-origin water (AW) and colder, polar origin water (PW) advecting southward in the East Greenland Current (EGC) influences the heat content of water entering Greenland's outlet glacial fjords. Here we use depth and temperature data derived from deep-diving seals to map out water mass variability across the continental shelf and to augment existing bathymetric products. We compare depths derived from the seal dives with the IBCAO Version 3 bathymetric database over the shelf and find differences up to 300 m near several large submarine canyons. In the vertical temperature structure, we find two dominant modes: a cold mode, with the typical AW/PW layering observed in the EGC, and a warm mode, where AW is present throughout the water column. The prevalence of these modes varies seasonally and spatially across the continental shelf, implying distinct AW pathways. In addition, we find that satellite sea surface temperatures (SST) correlate significantly with temperatures in the upper 50 m (R = 0.54), but this correlation decreases with depth (R = 0.22 at 200 m), and becomes insignificant below 250 m. Thus, care must be taken in using SST as a proxy for heat content, as AW mainly resides in these deeper layers. Sample Unit Level Copyright

Andersen, J. M., Wiersma, Y. F., Stenson, G. B., Hammill, M. O., Rosing-Asvid, A., and Skern-Maurizen, M. 2013. Habitat selection by hooded seals (Cystophora cristata) in the Northwest Atlantic Ocean. – ICES Journal of Marine Science, 70:173–185. We examined annual habitat use for 65 hooded seals (32 adult females, 17 adult males, and 16 juveniles) equipped with satellite relay data loggers (SRDLs) in spring or summer during five field seasons (2004–2008). A combined approach using first passage time (FPT) analysis and a generalized additive model (GAM) was applied to test for habitat selection, with a focus on environmental parameters of depth, slope, ice, sea surface temperature (SST), and chlorophyll. The models were run on adult males, adult females, and juveniles separately, and the results identified SST, depth, and chlorophyll as the most important factors influencing habitat selection across all categories. Furthermore, males and females preferred similar habitat conditions, but were separated geographically, and by depth, at various times of the year. Males appeared to be more localized in their habitat use patterns, focusing their search effort in areas of complex seabed relief such as Baffin Bay, Davis Strait, and the Flemish cap, while females concentrated their search effort along shelf areas (e.g. the Labrador shelf). These findings support our hypothesis that hooded seals prefer areas where topography and oceanographic processes create favourable foraging conditions.

Frie, A. K., Fagerheim, K-A., Hammill, M. O., Kapel, F. O., Lockyer, C., Stenson, G. B., Rosing-Asvid, A., and Svetochev, V. 2011. Error patterns in age estimation of harp seals (Pagophilus groenlandicus): results from a transatlantic, image-based, blind-reading experiment using known-age teeth. – ICES Journal of Marine Science, 68: 1942–1953. Blind readings of known-age samples are the ultimate quality control method for age estimates based on hard tissues. Unfortunately, this is often not feasible for many species because of the scarcity of known-age samples. Based on a unique collection of known-age teeth of harp seals (age range: 1–18 years), ageing errors were evaluated in relation to true age, reader experience, sex, and tooth format (images vs. originals). Bias was estimated by linear models fitted to deviations from true age, and precision was estimated as their residual standard error. Image-based blind readings of 98 tooth sections by 14 readers, representing different levels of experience, generally showed high accuracy and precision up to a seal age of ∼8 years, followed by an increasingly negative bias and increased variance. Separate analyses were therefore conducted for young seals (1–7 years) and older seals. For young seals, moderate associations were found between reader experience and levels of bias, precision, and proportions of correct readings. For older seals, only precision levels showed a significant association with reader experience. Minor effects of sex and tooth format are unlikely to affect these main patterns. Observed errors, even for highly experienced readers, may affect important age-related parameters, emphasizing the importance of known-age calibration of the output from all readers.

International audience; Recent studies have shown that the complementary analysis of mercury (Hg) concentrations and stable isotopic ratios of nitrogen (δ15N) and carbon (δ13C) can be useful for investigating the trophic influence on the Hg exposure and accumulation in marine top predators. In this study, we propose to evaluate the interspecies variability of Hg concentrations in phocids from polar areas and to compare Hg bioaccumulation between both hemispheres. Mercury concentrations, δ15N and δ13C were measured in fur from 85 individuals representing 7 phocidae species, a Ross seal (Ommatophoca rossii), Weddell seals (Leptonychotes weddellii), crabeater seals (Lobodon carcinophaga), harbour seals (Phoca vitulina), grey seals (Halichoerus grypus), ringed seals (Phoca hispida), and a bearded seal (Erignathus barbatus), from Greenland, Denmark and Antarctica. Our results showed a positive correlation between Hg concentrations and δ15N values among all individuals. Seals from the Northern ecosystems displayed greater Hg concentrations, δ15N and δ13C values than those from the Southern waters. Those geographical differences in Hg and stable isotopes values were likely due to higher environmental Hg concentrations and somewhat greater number of steps in Arctic food webs. Moreover, dissimilarities in feeding habits among species were shown through δ15N and δ13C analysis, resulting in an important interspecific variation in fur Hg concentrations. A trophic segregation was observed between crabeater seals and the other species, resulting from the very specific diet of krill of this species and leading to the lowest observed Hg concentrations.

It is crucial to examine the genetic diversity and structure of small, isolated populations, especially those at the edge of their distribution range, because they are vulnerable to stochastic processes if genetic diversity is low and the isolation level is high, and because such populations provide insight into the consequences of population declines in a broader conservation context. The harbour seal (Phoca vitulina) population in Svalbard is the world's northernmost P. vitulina population. Nothing is known about the genetic diversity, distinctiveness, or origin of this small, marginalized mammalian population. Thus, the present study investigated its genetic status in the context of nearby P. vitulina in Iceland, south-east Greenland, and northern Norway: this species is depleted/threatened in all of these regions. A total of 174 samples distributed between the four locations were analysed using 15 polymorphic microsatellites and variation in the displacement loop region (D-loop). Each of the four locations was a genetically distinct population. The Svalbard population was highly genetically distinct, had reduced genetic diversity, received limited gene flow, had a rather low effective population size, and showed an indication of having experienced a bottleneck resulting from a recent population decline. The significant heterozygote excess observed in the Svalbard sample might be attributed to the low effective population size, which could initiate future population inbreeding effects. This phenomenon has not been reported earlier from other P. vitulina populations, but if the Svalbard population is experiencing inbreeding, this could reduce its resilience to climate change, disease outbreaks, or other perturbations. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102, 420–439.

The recent rapid increase in mass loss from the Greenland ice sheet is primarily attributed to an acceleration of outlet glaciers. Oceanographic data obtained in summer 2008 show that subtropical waters that reside year-round in the shelf ocean off Greenland continuously enter a large glacial fjord in East Greenland and contribute to melting at the glacier terminus. The recent rapid increase in mass loss from the Greenland ice sheet1,2 is primarily attributed to an acceleration of outlet glaciers3,4,5. One possible cause of this acceleration is increased melting at the ice–ocean interface6,7, driven by the synchronous warming8,9,10 of subtropical waters offshore of Greenland. However, because of the lack of observations from Greenland’s glacial fjords and our limited understanding of their dynamics, this hypothesis is largely untested. Here we present oceanographic data collected in Sermilik Fjord, East Greenland, by ship in summer 2008 and from moorings. Our data reveal the presence of subtropical waters throughout the fjord. These waters are continuously replenished through a wind-driven exchange with the shelf, where they are present all year. The temperature and renewal of these waters indicate that they currently cause enhanced submarine melting at the glacier terminus. Key controls on the melting rate are the volume and properties of the subtropical waters on the shelf, and the patterns of along-shore winds, suggesting that the glaciers’ acceleration has been triggered by a combination of atmospheric and oceanic changes. Our measurements provide evidence for a rapid advective pathway for the transmission of oceanic variability to the ice-sheet margins.

We estimated demographic parameters and current harvest risks for a population of polar bears (Ursus maritimus Phipps) inhabiting northern Smith Sound and Kane Basin, Canada and Greenland. Our demographic analysis included a detailed assessment of age- and sex-specific survival and recruitment from 141 marked polar bears, using information contained within the standing age distribution of captures and mark-recapture analysis. Total survival rates $$ (\ifmmode\expandafter\bar\else\expandafter\=\fi{x} \pm 1\;{\text{SE}}) $$ for females were: 0.374 ± 0.180 (cubs), 0.686 ± 0.157 (ages 1–4), and 0.967 ± 0.043 (ages 5+). Mean litter size was 1.67 ± 0.08 cubs. Females did not reproduce until at least age 6, which is late compared to other populations of polar bears. The model-averaged, mark–recapture estimate of mean abundance (±1 SE) for years 1994–1997 was 164 ± 35 bears. We incorporated demographic parameters and their variances into a harvest risk analysis (i.e., a stochastic, harvested population viability analysis, PVA). Results suggest that polar bears in the region were severely over-harvested during the mark–recapture interval (1992–1997). The current status of the population is unknown.

The present pilot study examined emerging per- and polyfluoroalkyl substances (PFASs), i.e., a suite of short chain perfluoroalkyl acids (PFAAs), PFAA precursors and replacement chemicals, and legacy PFASs (long chain length PFAAs) in livers from ringed seals, polar bears and, for the first time, killer whales from East Greenland collected in 2012-2013. Among the emerging PFASs, perfluorobutanesulfonic acid (PFBS) and F-53B (a chlorinated polyfluorinated ether sulfonic acid) were detected in Arctic wildlife, albeit at concentrations approximately four orders of magnitude lower compared to perfluorooctanesulfonic acid (PFOS). PFOS was positively correlated with F-53B, but not PFBS in all three species. A total of 17 PFASs were detected in killer whales, including in a mother-fetus pair, demonstrating maternal transfer. ∑PFAS concentrations in killer whales (269 ± 90 ng/g) were comparable to concentrations found in ringed seals (138 ± 7 ng/g), however, an order of magnitude lower compared to concentrations found in polar bear livers (2336 ± 263 ng/g). Patterns of long chain PFAAs in killer whales differed from the pattern in ringed seals and polar bears. Of the monitored PFAA precursors, only perfluorooctanesulfonamide (FOSA) was detected in all three species, and FOSA/PFOS ratios and isomer patterns indicated that killer whales have a potential lower metabolic capacity to degrade FOSA compared to polar bears and ringed seals.

This work was funded through the Atlantic Seal Research Programme, International Governance Programme (GBS and MOH), the Greenland Institute of Natural Resources (ARA), and a CFI grant to YFW. The authors also acknowledge the support of the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland) in the completion of this study. MASTS is funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions (LB). Many pinniped species perform a specific dive type, referred to as a ‘drift dive’, where they drift passively through the water column. This dive type has been suggested to function as a resting/sleeping or food processing dive, and can be used as an indication of feeding success by calculating the daily change in vertical drift rates over time, which reflects the relative fluctuations in buoyancy of the animal as the proportion of lipids in the body change. Northwest Atlantic hooded seals perform drift dives at regular intervals throughout their annual migration across the Northwest Atlantic Ocean. We found that the daily change in drift rate varied with geographic location and the time of year and that this differed between sexes. Positive changes in buoyancy (reflecting increased lipid stores) were evident throughout their migration range and although overlapping somewhat, they were not statistically associated with high use areas as indicated by First Passage Time (FPT). Differences in the seasonal fluctuations of buoyancy between males and females suggest that they experience a difference in patterns of energy gain and loss during winter and spring, associated with breeding. The fluctuations in buoyancy around the moulting period were similar between sexes. Publisher PDF

Within their circumpolar range, polar bears (Ursus maritimus) are not subject to absolute barriers. However, physiographic features do cause discontinuities in their movements. These discontinuities in distribution can be used to delineate population units. Based on satellite telemetry of the movements of female polar bears carried out in 1989- 1998, we used cluster analysis to identify 6 regions within the Canadian and western Greenland Arctic in which move- ments appear to be restricted enough to identify distinct populations. These regions generally correspond to manage - ment units that have been previously identified as Viscount Melville Sound, Lancaster Sound, Norwegian Bay, Kane Basin, Baffin Bay, and Davis Strait. A north-south substructure was identified for the Baffin Bay population, but it was weaker than the structure identified for the 6 primary units. The 6 units were consistent with genetic information, ex - cept for the Baffin Bay - Kane Basin separation, and with mark-recapture observations and the traditional knowledge of Inuit hunters. Only 2 of 65 bears that provided telemetry information for more than 1 year were classified in differ - ent populations in different years. However, annual rates of exchange, measured as the percentage of locations outside the population boundary, ranged from 0.4 to 8.9%. Analysis of mark-recapture movements indicated no difference in large-scale movements between the sexes or long-term movements with age. Although our validation criteria for demo- graphic closure were satisfied, the observed rates of exchange between adjacent populations suggest that population dynamics in adjacent populations may not be completely independent. Resume : Il n'existe pas de barrieres absolues aux mouvements des Ours blancs (Ursus maritimus) dans leur reparti- tion ciricumpolaire; cependant, des caracteristiques physiographiques peuvent entrainer des discontinuites dans leurs de- placements, discontinuites qui peuvent servir a etablir les limites d'unites demographiques. Grâce a la telemetrie par satellite, nous avons suivi les deplacements de femelles de 1989 a 1998; une analyse de groupements a permis d'identifier 6 regions arctiques au Canada et dans l'ouest du Groenland ou les deplacements semblent suffisamment restreints pour identifier des populations distinctes. Ces regions correspondent aux unites d'amenagement identifiees precedemment : le detroit de Melville, le detroit de Lancaster, la baie Norwegian, le bassin de Kane, la baie de Baffin et le detroit de Davis. La population de la baie de Baffin compte en outre une sous-structure nord-sud qui est cepen- dant plus faible que la structure qui delimite les 6 unites primaires. Les 6 unites correspondent aux informations gene - tiques, sauf en ce qui concerne la separation baie de Baffin - bassin de Kane, et s'accordent aussi avec les donnees de capture-recapture et avec les connaissances traditionnelles de chasse des Inuit. Seulement 2 ours sur les 65 qui ont fourni des donnees telemetriques pendant plus de 1 annee ont ete classifies dans des populations differentes d'une annee a l'autre. Cependant, les taux annuels d'echange equivalant au pourcentage de sites au-dela des limites de la po - pulation, se situaient entre 0,4 et 8,9 %. L'analyse des donnees capture-recapture n'indique pas de difference entre les mâles et les femelles quant a leurs deplacements sur une grande echelle, ni entre individus d'âges differents quant a leurs deplacements a long terme. Bien que ce systeme reponde aux criteres de validation d'un systeme ferme, les taux d'echanges observes entre les populations adjacentes indiquent que la dynamique de chacune de ces populations n'est probablement pas totalement independante. (Traduit par la Redaction) Taylor et al. 709

We examined shelter and maternity dens used by 97 adult female polar bears (Ursus maritimus) in relation to conditions of sea ice. Obligate use of maternity dens for pregnancy, birth, and lactation varied little with latitude or area. In contrast, timing of facultative use of shelter dens switched from autumn in the southern area ( ,708N) to winter in the northern area (.758N). For the southern area, 13 of 16 female polar bears used shelter dens in autumn versus winter (median dates of entry and exit, 11 September and 2 November; total 5 56 days), whereas in the northern area, 11 of 17 bears used shelter dens in winter versus autumn (median dates, 24 December and 2 March; total 5 65 days). Difference in facultative use of shelter dens was associated with conditions of sea ice. Southern regions had no sea ice when polar bears used shelter dens. In contrast, northern areas had more constant ice conditions that included presence of ice throughout the year. Southern regions seem to have greater primary productivity and more seals as a result of a pronounced seasonal cycle of annual ice. Polar bears in northern areas responded to the more constant ice conditions and less productive environment with use of shelter dens during the period with lowest seal accessibility.