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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.

Since the late 1700s, reports of polar bears (Ursus maritimus) using tools (i.e., pieces of ice or stones) to kill walruses (Odobenus rosmarus) have been passed on verbally to explorers and naturalists by their Inuit guides, based on local traditional ecological knowledge (TEK) as well as accounts of direct observations or interpretations of tracks in the snow made by the Inuit hunters who reported them. To assess the possibility that polar bears may occasionally use tools to hunt walruses in the wild, we summarize 1) observations described to early explorers and naturalists by Inuit hunters about polar bears using tools, 2) more recent documentation in the literature from Inuit hunters and scientists, and 3) recent observations of a polar bear in a zoo spontaneously using tools to access a novel food source. These observations and previously published experiments on brown bears (Ursus arctos) confirm that, in captivity, polar and brown bears are both capable of conceptualizing the use of a tool to obtain a food source that would otherwise not be accessible. Based on the information from all our sources, this may occasionally also have been the case in the wild. We suggest that possible tool use by polar bears in the wild is infrequent and mainly limited to hunting walruses because of their large size, difficulty to kill, and their possession of potentially lethal weapons for both their own defense and the direct attack of a predator.

Kane Basin (KB) is one of the world's most northerly polar bear (Ursus maritimus) subpopulations, where bears have historically inhabited a mix of thick multiyear and annual sea ice year-round. Currently, KB is transitioning to a seasonally ice-free region because of climate change. This ecological shift has been hypothesized to benefit polar bears in the near-term due to thinner ice with increased biological production, although this has not been demonstrated empirically. We assess sea-ice changes in KB together with changes in polar bear movements, seasonal ranges, body condition, and reproductive metrics obtained from capture-recapture (physical and genetic) and satellite telemetry studies during two study periods (1993-1997 and 2012-2016). The annual cycle of sea-ice habitat in KB shifted from a year-round ice platform (~50% coverage in summer) in the 1990s to nearly complete melt-out in summer (

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.

Loss of sea ice brought on by climate change affects polar bear Ursus maritimus access to prey. Here we investigated variation in feeding habits of the Baffin Bay (BB) polar bear subpopulation in relation to sea ice, habitat use, season, and demography using hair carbon (δ13C), nitrogen (δ15N), and sulfur (δ34S) stable isotope values and total mercury (THg) concentrations as ecological tracers. We analyzed hair samples from BB polar bears (n = 131) of all age and sex classes live-captured in West Greenland during the spring in 2009-2013. BB polar bears occupied a narrow isotopic space, suggesting limited variation in carbon sources and trophic position within the subpopulation. THg concentrations (median ± SE: 5.1 ± 0.2, range: 0.3-12.5 µg g-1 dry weight, DW) were related to age class, and nearly half exceeded the suggested threshold for neurological effects in polar bears at 5.4 µg g-1 DW. Although distinct coastal and offshore space-use strategies have been reported for BB polar bears, our results suggest that both strategies lead to similar carbon sources and trophic positions. We found seasonal variation in δ13C and δ34S across both space-use strategies, with δ34S suggesting that all BB polar bears may prey on a higher proportion of benthic-feeding bearded seals Erignathus barbatus in late summer relative to spring. Despite wide fluctuations in inter-annual sea ice conditions and differences in space-use strategies among individuals, stable isotope values and THg concentrations suggested limited variation in feeding habits among BB polar bears. The variation of habitat tracers (δ13C and δ34S) was related to season, whereas trophic tracer (δ15N and THg) variation was driven by demographic group. The specialized BB polar bear diet suggests limited feeding plasticity under continued climate warming.

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.

The walrus (Odobenus rosmarus) is classified as a focal ecosystem component of the Arctic, defined as a biological element that is considered central to the functioning of an ecosystem, is of major importance to Arctic residents and/or is likely to be a good proxy for short- and long-term changes in the environment. The Arctic is undergoing large-scale environmental changes due to rapid global warming, including a marked reduction of sea ice in several areas inhabited by walruses. This chapter reviews how walruses already have been affected by global warming, or likely will be in the future. Specifically, we review the effects on walruses of projected changes in sea ice cover, marine productivity, ocean acidification, predation, pathogens and ultraviolet radiation, whereas changes in human activity patterns are discussed elsewhere in this volume. We find that, while the Pacific walrus seems to experience negative effects of warming and decrease in sea ice, the Atlantic walruses may be less affected; also in comparison to other ice-associated pinnipeds. Hence, we concur with previous assessments that the walrus is likely to survive into the future; at least in areas where human disturbance is minimal, and suitable terrestrial haul-outs are close enough to their feeding grounds.

Chapter 3 is a comprehensive overview of the ecology and behaviour of Atlantic walrus (Odobenus rosmarus rosmarus). We describe the classification, distribution, habitat requirements, morphology, reproduction, vital parameters, general behaviour, foraging and energy requirements, natural predators, diseases and parasites.

This chapter is a comprehensive overview of anthropogenic impacts on Atlantic walrus (Odobenus rosmarus rosmarus). We describe how Atlantic walruses are affected by hunting, fishery, anthropogenic noise (ships, aircraft), industrial activities (marine oil and gas exploration and exploitation, and mining), pollution and tourism.

Changes in sea-ice dynamics are affecting polar bears Ursus maritimus across their circumpolar range, which highlights the importance of periodic demographic assessments to inform management and conservation. We used genetic mark-recapture-recovery to derive estimates of abundance and survival for the Baffin Bay (BB) polar bear subpopulation—the first time this method has been used successfully for this species. Genetic data from tissue samples we collected via biopsy darting were combined with historical physical capture and harvest recovery data. The combined data set consisted of 1410 genetic samples (2011-2013), 914 physical captures (1993-1995, 1997), and 234 harvest returns of marked bears (1993-2013). The estimate of mean subpopulation abundance was 2826 (95% CI = 2284-3367) in 2012-2013. Estimates of annual survival (mean ± SE) were 0.90 ± 0.05 and 0.78 ± 0.06 for females and males age ≥2 yr, respectively. The proportion of total mortality of adult females and males that was attributed to legal harvest was 0.16 ± 0.05 and 0.26 ± 0.06, respectively. Remote sensing sea-ice data, telemetry data, and spatial distribution of onshore sampling indicated that polar bears were more likely to use offshore sea-ice habitat during the 1990s sampling period compared to the 2010s. Furthermore, in the 1990s, sampling of deep fjords and inland areas was limited, and no offshore sampling occurred in either time period, which precluded comparisons of abundance between the 1993-1997 and 2011-2013 study periods. Our findings demonstrate that genetic sampling can be a practical method for demographic assessment of polar bears over large spatial and temporal scales.

Climate change has broad ecological implications for species that rely on sensitive habitats. For some top predators, loss of habitat is expected to lead to cascading behavioral, nutritional, and reproductive changes that ultimately accelerate population declines. In the case of the polar bear (Ursus maritimus), declining Arctic sea ice reduces access to prey and lengthens seasonal fasting periods. We used a novel combination of physical capture, biopsy darting, and visual aerial observation data to project reproductive performance for polar bears by linking sea ice loss to changes in habitat use, body condition (i.e., fatness), and cub production. Satellite telemetry data from 43 (1991–1997) and 38 (2009–2015) adult female polar bears in the Baffin Bay subpopulation showed that bears now spend an additional 30 d on land (90 d in total) in the 2000s compared to the 1990s, a change closely correlated with changes in spring sea ice breakup and fall sea ice formation. Body condition declined for all sex, age, and reproductive classes and was positively correlated with sea ice availability in the current and previous year. Furthermore, cub litter size was positively correlated with maternal condition and spring breakup date (i.e., later breakup leading to larger litters), and negatively correlated with the duration of the ice‐free period (i.e., longer ice‐free periods leading to smaller litters). Based on these relationships, we projected reproductive performance three polar bear generations into the future (approximately 35 yr). Results indicate that two‐cub litters, previously the norm, could largely disappear from Baffin Bay as sea ice loss continues. Our findings demonstrate how concurrent analysis of multiple data types collected over long periods from polar bears can provide a mechanistic understanding of the ecological implications of climate change. This information is needed for long‐term conservation planning, which includes quantitative harvest risk assessments that incorporate estimated or assumed trends in future environmental carrying capacity.

Loss of Arctic sea ice has implications for the distribution and population structure of ice-dependent species such as polar bears Ursus maritimus. We used remotely sensed sea-ice concentration data for Baffin Bay, Canada, and satellite telemetry for adult female polar bears in the 1990s (n = 43) and 2000s (n = 38) to assess whether sea-ice habitat changes have influenced movements and habitat selection. Both the timing and availability of sea-ice habitat changed significantly between the 1990s and 2000s. Mean sea-ice concentration in June-October declined from 22 to 12%. Spring sea-ice retreat occurred 2 wk earlier and fall sea-ice advance 2 wk later in the 2000s. These changes translated directly to changes in habitat use by polar bears. In the 2000s, bears used significantly lower sea-ice concentrations in winter and spring. Also, bears were significantly closer to land in all months, except at the end of spring breakup when they remained on offshore sea ice as long as possible, likely to maximize foraging time prior to coming on land where they are largely food deprived. The presence of summer offshore sea ice facilitated broad movement of bears in the 1990s; however, this ice disappeared in the 2000s and resulted in significant declines in monthly movement rates. In the 2000s, adult females selected for lower sea-ice concentrations if they facilitated access to the continental shelf (

Identifying genetically different groups of animals, occupying specific geographical areas, is a prerequisite for conservation and management priorities. In the present study, the genetic structure of Atlantic walruses (Odobenus rosmarus rosmarus) occupying the Pechora Sea (PEC) in the western Russian Arctic, including walruses from Svalbard-Franz Josef Land (SVA-FJL) and East Greenland (EGR) regions, was investigated using 14 microsatellites (N = 159) and mtDNA sequences (N = 212). Bayesian-based clustering analysis identified two clusters: EGR and the other Northeast Atlantic areas. Pairwise F-ST analyses based on microsatellites revealed low but significant genetic differences between walruses from the PEC and SVA-FJL groups, which was supported by mtDNA analysis. F-ST was not significant for all sampling years, indicating a temporal effect or male-biased gene flow. Extended Bayesian Skyline Plots suggested a constant female subpopulation size (N-ef) for EGR and an increase for the SVA-FJL and PEC groups that commenced around 40-30 Kyr ago, indicating different demographic histories for walruses in the EGR. Further, the evolutionary phylogenetic relationship between Atlantic and Pacific walruses (O. r. divergence), based on mtDNA sequences, showed a monophyletic Atlantic clade, suggesting that Atlantic and Pacific walruses diverged similar to 949 Kyr. The principal finding suggests that PEC walruses show low, but significant genetic distinction from walruses in SVA-FJL and should be managed conservatively, as a separate, small population.

The phenology and habitat selection of polar bear (Ursus maritimus) maternity dens may shift over time in response to changing environmental conditions. We compared maternity den phenology and habitat characteristics using satellite telemetry data from adult female polar bears from the Baffin Bay (BB) (n = 16 dens; 2009–2015) and Kane Basin (KB) subpopulations (n = 3 dens; 2012–2015) to previously published maternity den data from 1991 to 1997 (BB n = 8 dens; KB n = 3 dens). BB maternity denning duration decreased from a mean of 194.1 days (SD = 21.0 days, n = 8) in the 1990s, to a mean of 167.1 days (SD = 27.6 days, n = 16; p = 0.017) in the 2000s. Delayed den entry accounted for shorter denning durations (1990s entry date $$\bar{x}$$ = 7 September; 2000s entry date $$\bar{x}$$ = 5 October; p = 0.018). For dens habitat characteristics of which could be measured, BB maternity dens in the 2000s occurred at higher elevations ( $$\bar{x}$$ = 707.0 m, SD = 284.9 m, n = 15; p = 0.003) and greater slopes ( $$\bar{x}$$ = 23.1°, SD = 7.4°; p = 0.003) than the 1990s (elevation $$\bar{x}$$ ± SD = 351.3 ± 194.5 m, n = 8; slope $$\bar{x}$$ ± SD = 11.9 ± 6.4°). Aspect also significantly differed between the 1990s ( $$\bar{x}$$ = 51.3°) and 2000s BB maternity dens ( $$\bar{x}$$ = 199.7°; Watson’s U 2 p = 0.042). KB dens were not statistically compared due to low sample size (n = 3 dens in both periods). Shifts in sea ice phenology and snow availability may explain the observed changes.

Knowledge of subpopulation identity including substructure is a prerequisite for sound management of polar bears (Ursus maritimus). It is not known whether the present catch of polar bears in the East Greenland subpopulation (EG) is sustainable. We used the Mean Measure of Divergence (MMD) to examine geographical variation in non-metrical traits from 1414 polar bear (Ursus maritimus) skulls collected in East Greenland (EG), Svalbard (SVA), Franz Josef Land (FJL), Davis Strait (DS), Baffin Bay (BB), and Kane Basin (KB), between 1830 and 2013. We focused on East Greenland with the goal of examining substructuring in the subpopulation. We did not find significant differences among samples across four areas of the EG subpopulation (i.e., offshore Fram Strait, NE, SE, and SW Greenland) using data from 1830 to 1983. Our analyses did not lend support to substructuring. However, we draw our conclusions with caution because skulls were sampled over a long time period and had low power due to small sample sizes. Also, comparisons were limited to pre-1980s skulls. The decrease in sea ice in EG since the 1990s due to climate change may have led to substructuring not detected with MMD. This study contributes to the current efforts by Greenland authorities to quantify connectivity of polar bears between southeast and northeast Greenland which is important information for the evaluation of the sustainability of the catch of bears from the EG subpopulation.

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/).

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.

Polar bears (Ursus maritimus) are among the world's highest trophic level marine predators and as such have some of the highest tissue concentrations of organohalogen contaminants (OHCs) among Arctic biota. In this paper we present the results of a three decade (1983–2013) risk assessment of OHC exposure and effects on reproduction, immunity, and cancer (genotoxicity) in polar bears from Central East Greenland. Risk of adverse effects are evaluated using a risk quotient (RQ) approach with derivation from measured OHC concentrations in polar bear tissue and critical body residues (CBR) extrapolated for polar bears using physiologically-based pharmacokinetic modelling (PBPK). The additive RQs for all OHCs in polar bears were above the threshold for all effect categories (RQ > 1) in every year, suggesting this population has been at significant and continuous risk of contaminant-mediated effects for over three decades. RQs peaked in 1983 (RQ > 58) and again in 2013 (RQ > 50) after a period of decline. These trends follow ΣPCB levels during that time, and contributed almost all of the risk to immune, reproductive, and carcinogenic effects (71–99% of total RQ). The recent spike in RQs suggests a major shift in polar bear contaminant exposure from climate related changes in food composition and hereby the increased risk of adverse health effects. In the context of lifetime exposure ΣPCB and PFOS levels showed the interactive importance of year of birth, age, and emission history. In conclusion, the results indicate that East Greenland polar bears have been exposed to OHC levels over the period of 1983–2013 that potentially and continuously affected individual and theoretically also population health, with a peaking risk in the more recent years. Keywords: Polar bear, Ursus maritimus, Organohalogen contaminants, PBPK modelling, Critical body residue, Immune suppression, Reproductive toxicity, Risk quotient

Vitamins A (retinol) and E (α-tocopherol) are dietary vitamins, essential for, e.g., growth and development, reproduction, and immune function. Persistent organic pollutants (POPs) have been found to be related to vitamin A and E metabolism. However, few investigations have been published on this health issue in polar bears (Ursus maritimus). The aim of this study was thus to provide reference values for concentrations of vitamin A in liver, kidney cortex, and whole blood and vitamin E in kidney cortex and whole blood from 166 East Greenland polar bears, as well as to assess the relationship between POPs and vitamin concentrations. In addition, vitamin concentrations were analyzed for temporal trends (1994–2008). Results showed vitamin A in liver to be higher in adult bears and the concentrations of vitamin E in kidney and blood to likewise be generally higher in adult bears. In addition, all analyzed contaminant groups were correlated with at least one of the vitamin parameters, predominantly in a negative way. Finally, vitamin A liver concentrations as well as concentration of vitamin E in kidney and blood showed a temporal increase. Together, these results add to the weight of evidence that POPs could be disrupting polar bear vitamin status. However, while the observed temporal increases in vitamin concentrations were likely POP related, the question remains as to whether they stem from influence of contaminants only or also, e.g., changes in prey species. Further studies are needed to tease apart the causes underlying these changes in vitamin concentrations.

Arctic marine mammals (AMMs) are icons of climate change, largely because of their close association with sea ice. However, neither a circumpolar assessment of AMM status nor a standardized metric of sea ice habitat change is available. We summarized available data on abundance and trend for each AMM species and recognized subpopulation. We also examined species diversity, the extent of human use, and temporal trends in sea ice habitat for 12 regions of the Arctic by calculating the dates of spring sea ice retreat and fall sea ice advance from satellite data (1979–2013). Estimates of AMM abundance varied greatly in quality, and few studies were long enough for trend analysis. Of the AMM subpopulations, 78% (61 of 78) are legally harvested for subsistence purposes. Changes in sea ice phenology have been profound. In all regions except the Bering Sea, the duration of the summer (i.e., reduced ice) period increased by 5–10 weeks and by >20 weeks in the Barents Sea between 1979 and 2013. In light of generally poor data, the importance of human use, and forecasted environmental changes in the 21st century, we recommend the following for effective AMM conservation: maintain and improve comanagement by local, federal, and international partners; recognize spatial and temporal variability in AMM subpopulation response to climate change; implement monitoring programs with clear goals; mitigate cumulative impacts of increased human activity; and recognize the limits of current protected species legislation.

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.

Background Satellite telemetry studies provide information that is critical to the conservation and management of species affected by ecological change. Here we report on the performance and retention of two types (SPOT-227 and SPOT-305A) of ear-mounted Argos-linked satellite transmitters (i.e., platform transmitter terminal, or PTT) deployed on free-ranging polar bears in Eastern Greenland, Baffin Bay, Kane Basin, the southern Beaufort Sea, and the Chukchi Sea during 2007–2013. Results Transmissions from 142 out of 145 PTTs deployed on polar bears were received for an average of 69.3 days. The average functional longevity, defined as the number of days they transmitted while still attached to polar bears, for SPOT-227 was 56.8 days and for SPOT-305A was 48.6 days. Thirty-four of the 142 (24%) PTTs showed signs of being detached before they stopped transmitting, indicating that tag loss was an important aspect of tag failure. Furthermore, 10 of 26 (38%) bears that were re-observed following application of a PTT had a split ear pinna, suggesting that some transmitters were detached by force. All six PTTs that were still on bears upon recapture had lost the antenna, which indicates that antenna breakage was a significant contributor to PTT failure. Finally, only nine of the 142 (6%) PTTs—three of which were still attached to bears—had a final voltage reading close to the value indicating battery exhaustion. This suggests that battery exhaustion was not a major factor in tag performance. Conclusions The average functional longevity of approximately 2 months for ear-mounted PTTs (this study) is poor compared to PTT collars fitted to adult female polar bears, which can last for several years. Early failure of the ear-mounted PTTs appeared to be caused primarily by detachment from the ear or antenna breakage. We suggest that much smaller and lighter ear-mounted transmitters are necessary to reduce the risk of tissue irritation, tissue damage, and tag detachment, and with a more robust antenna design. Our results are applicable to other tag types (e.g., iridium and VHF systems) and to research on other large mammals that cannot wear radio collars.

Legacy organochlorine contaminants were determined in adipose tissues from 294 polar bears (Ursus maritimus) sampled in East Greenland in 23 of the 28 years between 1983 and 2010. Of 19 major legacy contaminants and congeners (ΣPCB, 4 PCB congeners (CB153, 180, 170/190), ΣDDT, p,p′-DDE, p,p′ -DDD and p,p′-DDT, α- and β-hexachlorocyclohexane (HCH), HCB, octachlorostyrene, dieldrin, oxychlordane, cis- and trans-chlordane, cis- and trans-nonachlor, heptachlor epoxide and BB-153), 18 showed statistically significant average yearly declines of −4.4% (range: −2.0 to −10.8%/year) among subadult polar bears (i.e. females

International audience; The Arctic is facing major environmental changes impacting marine biodiversity and ecosystem functioning. One way of assessing the responses of an ecosystem to these changes is to quantitatively study food web dynamics. Here, we used stable isotope (δ15N and δ13C) analyses of 39 Arctic marine species to investigate trophic relationships and isotopic niches of the West Greenland food web in 2000–2004. The lowest δ15N values were found for suspension feeding blue mussel (Mytilus edulis; 6.1 ‰) and the highest for polar bear (Ursus maritimus; 20.2 ‰). For δ13C, copepods (Calanus spp.) had the lowest values (−20.4 ‰) and snow crab (Chionoecetes opilio) the highest values (−15.8 ‰). Our results show that the three trophic enrichment factor (TEF) approaches used to quantify species trophic positions (fixed TEF of 3.8 and 3.4 ‰ or scaled TEF) did not generally affect trophic modelling and provided similar conclusions. Overall, the findings in this study are in good agreement with previous investigations of other Arctic marine ecosystems. Interestingly, we found little overlap of core isotopic niches used by the four investigated functional groups (mammals, seabirds, fish and invertebrates), except for seabirds and fish where an overlap of 24 % was found. These results provide new insights into species and functional group interactions, as well as into the food web structure and ecosystem functioning of an important Arctic region that can be used as a template to guide future modelling of carbon, energy and contaminant flow in the region.

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

Several perfluoroalkyl carboxylates (PFCAs) and perfluoroalkyl sulfonates (PFSAs) of varying chain length are bioaccumulative in biota. However, wildlife reports have focused on liver and with very little examination of other tissues, and thus there is a limited understanding of their distribution and potential effects in the mammalian body. In the present study, the comparative accumulation of C(6) to C(15) PFCAs, C(4), C(6), C(8) and C(10) PFSAs, and select precursors were examined in the liver, blood, muscle, adipose, and brain of 20 polar bears (Ursus maritimus) from Scoresby Sound, Central East Greenland. Overall, PFSA and PFCA concentrations were highest in liver followed by bloodbrainmuscle ≈ adipose. Liver and blood samples contained proportionally more of the shorter/medium chain length (C(6) to C(11)) PFCAs, whereas adipose and brain samples were dominated by longer chain (C(13) to C(15)) PFCAs. PFCAs with lower lipophilicities accumulated more in the liver, whereas the brain accumulated PFCAs with higher lipophilicities. The concentration ratios (±SE) between perfluorooctane sulfonate and its precursor perfluorooctane sulfonamide varied among tissues from 9 (±1):1 (muscle) to 36 (±7):1 (liver). PFCA and PFSA patterns in polar bears indicate that the pharmacokinetics of these compounds are to some extent tissue-specific, and are the result of several factors that may include differing protein interactions throughout the body.

The multivariate relationship between hair cortisol, whole blood thyroid hormones, and the complexmixtures of organohalogen contaminant (OHC) levels measured in subcutaneous adipose of 23 EastGreenland polar bears (eight males and 15 females, all sampled between the years 1999 and 2001) wasanalyzed using projection to latent structure (PLS) regression modeling. In the resulting PLS model,most important variables with a negative influence on cortisol levels were particularly BDE-99, but alsoCB-180, -201, BDE-153, and CB-170/190. The most important variables with a positive influence oncortisol were CB-66/95, a -HCH, TT3, as well as heptachlor epoxide, dieldrin, BDE-47, p,p 0 -DDD.Although statistical modeling does not necessarily fully explain biological cause–effect relationships,relationships indicate that (1) the hypothalamic–pituitary–adrenal (HPA) axis in East Greenland polarbears is likely to be affected by OHC-contaminants and (2) the association between OHCs and cortisolmay be linked with the hypothalamus–pituitary–thyroid (HPT) axis.& 2012 Elsevier Inc. All rights reserved.

One of the primary mechanisms by which sea ice loss is expected to affect polar bears is via reduced body condition and growth resulting from reduced access to prey. To date, negative effects of sea ice loss have been documented for two of 19 recognized populations. Effects of sea ice loss on other polar bear populations that differ in harvest rate, population density, and/or feeding ecology have been assumed, but empirical support, especially quantitative data on population size, demography, and/or body condition spanning two or more decades, have been lacking. We examined trends in body condition metrics of captured bears and relationships with summertime ice concentration between 1977 and 2010 for the Baffin Bay (BB) and Davis Strait (DS) polar bear populations. Polar bears in these regions occupy areas with annual sea ice that has decreased markedly starting in the 1990s. Despite differences in harvest rate, population density, sea ice concentration, and prey base, polar bears in both populations exhibited positive relationships between body condition and summertime sea ice cover during the recent period of sea ice decline. Furthermore, females and cubs exhibited relationships with sea ice that were not apparent during the earlier period (1977–1990s) when sea ice loss did not occur. We suggest that declining body condition in BB may be a result of recent declines in sea ice habitat. In DS, high population density and/or sea ice loss, may be responsible for the declines in body condition.

A statistically robust method was applied to 83 time-series of mercury in Arctic biota from marine, freshwater and terrestrial ecosystems with the purpose of generating a 'meta-analysis' of temporal trend data collected over the past two to three decades, mostly under the auspices of the Arctic Monitoring and Assessment Program (AMAP). Sampling locations ranged from Alaska in the west to northern Scandinavia in the east. Information from recently published temporal trend studies was tabulated to supplement the results of the statistical analyses. No generally consistent trend was evident across tissues and species from the circumpolar Arctic during the last 30years or so. However, there was a clear west-to-east gradient in the occurrence of recent increasing Hg trends, with larger numbers and a higher proportion of biotic datasets in the Canadian and Greenland region of the Arctic showing significant increases than in the North Atlantic Arctic. Most of the increasing datasets were for marine species, especially marine mammals. A total of 16 (19%) out of the 83 time-series could be classified as "adequate", where adequate is defined as the number of actual monitoring years in a time-series being equal to or greater than the number of years of sampling required to detect a 5% annual change in Hg concentrations, with a significance level of P

Thyroid gland histology was examined in 20 organohalogen contaminant (OHC)-exposed East Greenland polar bears (Ursus maritimus). OHC concentrations measured in subcutaneous adipose tissue were between 3556 and 28,670 ng g−1 lw for polychlorinated biphenyls (ΣPCB51), 9 and 3403 ng g−1 lw for the sum of organochlorine pesticides (hexachlorocyclohexanes, hexachlorobenzene, chlordanes, dichlorodiphenyltrichloroethanes, and dieldrin), and from 21 to 130 ng g−1 lw for polybrominated diphenyl ethers. Histological examinations revealed that 12 of the bears (10 males aged 3–19 years and two females aged 4–7 years) had normal thyroid tissue while eight bears (40%) of varying ages and genders (three males aged 3–9 years and five females aged 4–25 years) showed clear histological lesions including parafollicular C-cell proliferation, nodular hyperplasia, and interstitial fibrosis. No significant differences were found in prevalence of thyroid gland lesions between males and females. Similarly, no marked difference wa...

The relative contribution of regional contamination versus dietary differences to geographic variation in polar bear (Ursus maritimus) contaminant levels is unknown. Dietary variation between Alaska, Canada, East Greenland, and Svalbard subpopulations was assessed by muscle nitrogen and carbon stable isotope (δ(15)N, δ(13)C) and adipose fatty acid (FA) signatures relative to their main prey (ringed seals). Western and southern Hudson Bay signatures were characterized by depleted δ(15)N and δ(13)C, lower proportions of C(20) and C(22) monounsaturated FAs and higher proportions of C(18) and longer chain polyunsaturated FAs. East Greenland and Svalbard signatures were reversed relative to Hudson Bay. Alaskan and Canadian Arctic signatures were intermediate. Between-subpopulation dietary differences predominated over interannual, seasonal, sex, or age variation. Among various brominated and chlorinated contaminants, diet signatures significantly explained variation in adipose levels of polybrominated diphenyl ether (PBDE) flame retardants (14-15%) and legacy PCBs (18-21%). However, dietary influence was contaminant class-specific, since only low or nonsignificant proportions of variation in organochlorine pesticide (e.g., chlordane) levels were explained by diet. Hudson Bay diet signatures were associated with lower PCB and PBDE levels, whereas East Greenland and Svalbard signatures were associated with higher levels. Understanding diet/food web factors is important to accurately interpret contaminant trends, particularly in a changing Arctic.

Defining sub-populations and determining migration rates between them is crucial for sound management and conservation. This also applies to the Atlantic walrus Odobenus rosmarus rosmarus, which is still exploited although population levels are low in some cases and information on abundance is lacking in others. Analysis of muscle and skin tissues using 11 nuclear microsatellite markers from a total of 297 Atlantic walruses from the Hudson Strait across W, NW and E Greenland to Svalbard and Franz Josef Land was undertaken to determine the number of sub-populations, their ancestral origin and the contemporary rates and directions of migration (gene flow) between the var- ious areas. The study indicated the existence of 5 sub-populations in the Hudson Strait, W Greenland, NW Greenland, E Greenland and Franz Josef Land-Svalbard. Identification of the Hudson Strait sub- population was novel; although differences between animals here and in W Greenland were small, they were statistically significant and indicated that walruses in the Hudson Strait could be a popu- lation source for walruses for the W Greenland sub-population. The direction of migration (Hudson Strait to W Greenland) suggested a genetic signal from a historical large-scale counter-clockwise perennial migration pattern in the Baffin Bay region. The study provides essential information on population sub-structuring that is a prerequisite for management of Atlantic walruses at sustainable levels.

Polar bears (Ursus maritimus) feed mainly on ringed seal (Phoca hispida) and consume large quantities of blubber and consequently have one of the highest tissue concentrations of organohalogen contaminants (OHCs) worldwide. In East Greenland, studies of OHC time trends and organ system health effects, including reproductive, were conducted during 1990-2006. However, it has been difficult to determine the nature of the effects induced by OHC exposures on wild caught polar bears using body burden data and associated changes in reproductive organs and systems. We therefore conducted a risk quotient (RQ) evaluation to more quantitatively evaluate the effect risk on reproduction (embryotoxicity and teratogenicity) based on the critical body residue (CBR) concept and using a physiologically-based pharmacokinetic (PBPK) model. We applied modelling approaches to PCBs, p,p'-DDE, dieldrin, oxychlordane, HCHs, HCB, PBDEs and PFOS in East Greenland polar bears based on known OHC pharmacokinetics and dynamics in laboratory rats (Rattus rattus). The results showed that subcutaneous adipose tissue concentrations of dieldrin (range: 79-1271 ng g(-1) lw) and PCBs (range: 4128-53,923 ng g(-1) lw) reported in bears in the year 1990 were in the range to elicit possible adverse health effects on reproduction in polar bears in East Greenland (all RQs > or = 1). Similar results were found for PCBs (range: 1928-17,376 ng g(-1) lw) and PFOS (range: 104-2840 ng g(-1) ww) in the year 2000 and for dieldrin (range: 43-640 ng g(-1) lw), PCBs (range: 3491-13,243 ng g(-1) lw) and PFOS (range: 1332-6160 ng g(-1) ww) in the year 2006. The concentrations of oxychlordane, DDTs, HCB and HCHs in polar bears resulted in RQs

The walrus (Odobenus rosmarus) is in some current systematic schemes divided into three subspecies: O. r. rosmarus in the North Atlantic, O. r. divergens in the North Pacific and O. r. laptevi in the Laptev Sea. These three subspecies have been described as differing in body size, but the taxonomic status of O. r. laptevi is disputed. The current study applies molecular and morphometric methods to assess the taxonomic status of O. r. laptevi and to analyse the systematic and phylogeographic relationships between the three purported walrus subspecies. Tusk length and tusk circumference were measured from the few skulls available of O. r. laptevi, and the obtained values were within the ranges reported for Pacific walruses. Thus, morphologically, subspecies status for O. r. laptevi is not supported according to the Amadon Mayr '75% rule'. Phylogenetic analyses and haplotype networks based on mitochondrial nucleotide sequence data of NADH dehydrogenase 1, 16S rRNA, cytochrome oxidase I and the D-loop of the control region of the historic O. r. laptevi bone material and contemporary O. r. rosmarus and O. r. divergens showed that the Laptev Sea walrus groups with individuals from the North Pacific. Thus, the mitochondrial sequence data do not support the recognition of three walrus subspecies as reciprocally monophyletic evolutionary units with independent evolutionary histories. Only O. r. rosmarus and O. r. divergens meet this criterion with the present sampling. Accordingly, we recommend that Odobenus r. laptevi be abandoned and the Laptev walrus instead be recognized as the westernmost population of the Pacific walrus, Odobenus r. divergens. However, further research is recommended to assess whether the Laptev walrus could be considered as a significant unit in terms of conservation and management, since it is unique in several ecological parameters. (Less)

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

In this article, we review the effects on polar bears of global warming that have already been observed, and try to evaluate what may happen to the polar bears in the future. Many researchers have predicted a wide range of impacts of climate change on polar bear demography and conditions. A predicted major reduction in sea ice habitat will reduce the availability of ice associated seals, the main prey of polar bears, and a loss and fragmentation of polar bear habitat will ultimately lead to large future reductions in most subpopulations. It is likely that polar bears will be lost from many areas where they are common today and also that the total population will change into a few more distinctly isolated populations.