Movements of female polar bears (<Emphasis Type="Italic">Ursus maritimus</Emphasis>) in the East Greenland pack ice

The movements of two adult female polar bears ( Ursus maritimus) in East Greenland and the Greenland Sea area were studied by use of satellite telemetry between the fall of 1994 and the summer of 1998. One female was tracked for 621 days, the other for 1,415 days. During this time the females used maternity dens on land. If denning periods on land were excluded, the two females used between 73% and 100% of the tracking time offshore where they were able to navigate in the dynamic pack ice and counteract the fast southward movement of the ice (up to 30 km/h) in the East Greenland Current. Mean monthly movement rates varied between 0.32 and 0.76km/h. Both bears had very large home ranges (242,000 and 468,000 km ²) within the dynamic pack ice of the Greenland Sea. The facts that the bears made extensive use of the offshore sea ice and that there is a marked reduction of the Greenland Sea ice call for a closer monitoring of the effects of this change on the East Greenland polar bear population.     

Polar bear stress hormone cortisol fluctuates with the North Atlantic Oscillation climate index

Polar bears are heavily dependent on sea ice for hunting sufficient prey to meet their energetic needs. When the bears are left fasting, it may cause a rise in the levels of the stress hormone cortisol. Cortisol is the major corticosteroid hormone in most mammals, including polar bears. Production and regulation of this stress hormone are vital for the body as it is part of a myriad of processes, including in relation to metabolism, growth, development, reproduction, and immune function. In the present study, we examined the correlation between East Greenland polar bear hair cortisol concentration (HCC), a matrix that reflects longer-term hormone levels, and the fluctuations of the North Atlantic Oscillation (NAO) index, a large-scale climate phenomenon applied as a proxy for sea ice extent in the Greenland Sea along the coast of East Greenland. In doing so, a significant positive correlation (r = 0.88; p = 0.0004) was found between polar bear hair cortisol and the NAO, explaining 77 % of the variation in HCC observed between years over the period 1989–2009. This result indicates that interannual fluctuations in climate and ice cover have a substantial influence on longer-term cortisol levels in East Greenland polar bears. Further research into the implications and consequences inherent in this correlation are recommended, preferably across multiple polar bear populations.     

Differences in growth,size and sexual dimorphism in skulls of East Greenland and Svalbard polar bears (<Emphasis Type="Italic">Ursus maritimus</Emphasis>)

Size, growth and sexual dimorphism of nine skull traits was studied in 300 East Greenland and 391 Svalbard polar bears (Ursus maritimus). Two traits were significantly larger in bears from East Greenland compared to Svalbard bears, and trait size was smaller after 1960 in five traits. For both localities and both age groups (subadult, adult), mean trait size values were higher in males than females (all: P < 0.05). Gompertz growth models showed trait size increasing with age in seven traits. Depending on the trait, males reached 95% asymptotic trait size at age 3–10, females at age 2–6. The females of both localities matured at approximately the same age, whereas the Svalbard males generally matured years later than their East Greenland peers. The differences found in the present study between the two polar bear subpopulations support the notion that East Greenland and Svalbard polar bears probably should be managed as separate units.     

Prevalence of antibodies against Toxoplasma gondii in polar bears (Ursus maritimus) from Svalbard and East Greenland

Serum samples from 419 polar bears (Ursus maritimus) from Svalbard and the Barents Sea (collected 1990-2000) and 108 polar bears from East Greenland (collected 1999-2004) were assayed for antibodies against Toxoplasma gondii using the modified agglutination test. Antibody prevalences were 3.6% among cubs dependent on their mothers and 21.4% among subadults and adults. Among subadults and adults there was an interaction between population and sex, with similar prevalences among females (Svalbard = 19.5%, Greenland = 18.0%), but a high frequency among Svalbard males (28.7%) as compared to Greenland males (5.8%). The pattern was also significant after correcting for differences in age distribution. The sex-population interaction term is believed to be connected to area- and sex-specific feeding ecology. The prevalences of antibodies against T. gondii in Svalbard and Greenland were high compared to previously reported findings in polar bears from Russian and Alaskan areas.     

Families in space: relatedness in the Barents Sea population of polar bears (Ursus maritimus)

The kin structure and dispersal pattern of polar bears ( Ursus maritimus ) of the Barents Sea was investigated during the spring mating season using two complementary approaches. First, individual genotypes based on the analyses of 27 microsatellite loci of 583 polar bears were related to field information gathered from 1146 bears in order to reconstruct the animals' pedigrees and to infer geographical distances between adult bears of different relatedness categories. According to the data, the median natal dispersal distance of the male animals was 52 km while that of the females was 93 km. Second, the relatedness of pairs of adult bears was estimated and correlated to the geographical distance between them. The female dyads had a much stronger kin structure than the male dyads. The 'pedigree approach' revealed a male kin structure which could not be detected using the 'relatedness approach'. This suggests that, on a broader scale, effective dispersal is slightly male biased. Despite fidelity to natal areas, male-mediated gene flow may nevertheless prevent genetic differentiation. Males might occasionally shift their home range which could therefore lead to a male-biased breeding dispersal. Our results showed that a nonterritorial species such as the polar bear that has a high dispersal potential, lives in a highly unstable environment and migrates seasonally is still able to exhibit a distinct kin structure during the mating season.     

Comparative fate of organohalogen contaminants in two top carnivores in Greenland: captive sledge dogs and wild polar bears

The limited knowledge and/or the inability to control physiological condition parameters that influence the fate of organohalogen contaminants (OHCs) has been the foremost confounding aspect in monitoring programs and health risk assessments of wild top predators in the Arctic such as the polar bear (Ursus maritimus). In the present comparative study, we used a potential surrogate Canoidea species for the East Greenland polar bear, the captive sledge dog (Canis familiaris), to investigate some factors that may influence the bioaccumulation and biotransformation of major chlorinated and brominated OHCs in adipose tissue and blood (plasma) of control (fed commercial pork fat) and exposed (fed West Greenland minke whale (Balaenoptera acutorostrata) blubber) adult female sledge dogs. Furthermore, we compared the patterns and concentrations of OHCs and their known or suggested hydroxylated (OH) metabolites (e.g., OH-PCBs) in sledge dogs with those in adipose tissue and blood (plasma) of East Greenland adult female polar bears, and blubber of their main prey species, the ringed seal (Pusa hispida). The two-year feeding regime conducted with sledge dogs led to marked differences in overall adipose tissue (and plasma) OHC residue accumulation between the control and exposed groups. Characteristic prey-to-predator OHC bioaccumulation dynamics for major PCB and PBDE congeners (patterns and concentrations) and biotransformation capacity with respect to PCB metabolite formation and OH-PCB retention distinguished, to some extent, captive sledge dogs and wild polar bears. Based on the present findings, we conclude that the use of surrogate species in toxicological investigations for species in the Canoidea family should be done with great caution, although they remain essential in the context of contaminants research with sensitive arctic top carnivore species such as the polar bear.     

Microsatellite analysis of population structure in Canadian polar bears

Attempts to study the genetic population structure of large mammals are often hampered by the low levels of genetic variation observed in these species. Polar bears have particularly low levels of genetic variation with the result that their genetic population structure has been intractable. We describe the use of eight hypervariable microsatellite loci to study the genetic relationships between four Canadian polar bear populations: the northern Beaufort Sea, southern Beaufort Sea, western Hudson Bay, and Davis Strait - Labrador Sea. These markers detected considerable genetic variation, with average heterozygosity near 60% within each population. Interpopulation differences in allele frequency distribution were significant between all pairs of populations, including two adjacent populations in the Beaufort Sea. Measures of genetic distance reflect the geographic distribution of populations, but also suggest patterns of gene flow which are not obvious from geography and may reflect movement patterns of these animals. Distribution of variation is sufficiently different between the Beaufort Sea populations and the two more eastern ones that the region of origin for a given sample can be predicted based on its expected genotype frequency using an assignment test. These data indicate that gene flow between local populations is restricted despite the long-distance seasonal movements undertaken by polar bears.     

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

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

Field Metabolic Rate and PCB Adipose Tissue Deposition Efficiency in East Greenland Polar Bears Derived from Contaminant Monitoring Data

Climate change will increasingly affect the natural habitat and diet of polar bears (Ursus maritimus). Understanding the energetic needs of polar bears is therefore important. We developed a theoretical method for estimating polar bear food consumption based on using the highly recalcitrant polychlorinated biphenyl (PCB) congener, 2,2′,4,4′,55-hexaCB (CB153) in bear adipose tissue as an indicator of food intake. By comparing the CB153 tissue concentrations in wild polar bears with estimates from a purposely designed individual-based model, we identified the possible combinations of field metabolic rates (FMR) and CB153 deposition efficiencies in East Greenland polar bears. Our simulations indicate that if 30% of the CB153 consumed by polar bear individuals were deposited into their adipose tissue, the corresponding FMR would be only two times the basal metabolic rate. In contrast, if the modelled CB153 deposition efficiency were 10%, adult polar bears would require six times more energy than that needed to cover basal metabolism. This is considerably higher than what has been assumed for polar bears in previous studies though it is similar to FMRs found in other marine mammals. An implication of this result is that even relatively small reductions in future feeding opportunities could impact the survival of East Greenland polar bears.     

Bear-hybrids: behaviour and phenotype

The behaviour of a female and a male bear hybrid (brown bear Ursus arctos x polar bear Ursus maritimus) was studied. Behavioural and morphological comparisons between the hybrids and the two parent-species were made.The observation period covered July to November 2007. Different objects were offered to the bears to evoke new patterns of behaviour. While manipulating the offered objects, both bears showed elements of behaviour also found in polar bears, though the male performed those to a larger extend. Not only in the way of manipulating objects but also in the male's type of stereotype parallels to polar bears can be drawn. Phenotypically both bears possess features of both species, brown bear and polar bear. Concerning it lighter colour of fur and the structure of hair the female hybrid resembles more a polar bear than the male. Summarising, the female appears phenotypically more like a polar bear, the male with respect to the behaviour.     

Brucella sp. antibodies in polar bears from Svalbard and the Barents Sea

A prevalence of 5.4% of anti-Brucella sp. antibodies was found in plasma samples from 297 polar bears (Ursus maritimus) from Svalbard and the Barents Sea. Plasma was tested by the classical brucellosis tests Slow Agglutination of Wright (SAW), EDTA modified SAW and Rose Bengal test, as well as by an indirect Protein A ELISA. Only samples classified as positive in all tests were regarded as containing anti-Brucella sp. antibodies. A significant west to east increase in the proportion of bears with anti-Brucella sp. antibodies was found, with 3.6% (n = 253) at Svalbard (Spitsbergen, Nordaustlandet, Edge?ya, Barents?ya and Hopen), and 15.9% (n = 44) in the central Barents Sea. Anti-Brucella sp. antibodies were previously found in ringed seals (Phoca hispida) and harp seals (Phoca groenlandica) from the same geographical areas. The ringed seal is an important prey species for the Svalbard polar bear population, and may thus be a source of brucellosis for the bears. There are no indications of reproductive disorders caused by Brucella sp. or other infectious agents in our study polar bear population. Potential impacts of Brucella sp. exposure on individuals or the population are unknown.     

Implications of the Circumpolar Genetic Structure of Polar Bears for Their Conservation in a Rapidly Warming Arctic

We provide an expansive analysis of polar bear (Ursus maritimus) circumpolar genetic variation during the last two decades of decline in their sea-ice habitat. We sought to evaluate whether their genetic diversity and structure have changed over this period of habitat decline, how their current genetic patterns compare with past patterns, and how genetic demography changed with ancient fluctuations in climate. Characterizing their circumpolar genetic structure using microsatellite data, we defined four clusters that largely correspond to current ecological and oceanographic factors: Eastern Polar Basin, Western Polar Basin, Canadian Archipelago and Southern Canada. We document evidence for recent (ca. last 1–3 generations) directional gene flow from Southern Canada and the Eastern Polar Basin towards the Canadian Archipelago, an area hypothesized to be a future refugium for polar bears as climate-induced habitat decline continues. Our data provide empirical evidence in support of this hypothesis. The direction of current gene flow differs from earlier patterns of gene flow in the Holocene. From analyses of mitochondrial DNA, the Canadian Archipelago cluster and the Barents Sea subpopulation within the Eastern Polar Basin cluster did not show signals of population expansion, suggesting these areas may have served also as past interglacial refugia. Mismatch analyses of mitochondrial DNA data from polar and the paraphyletic brown bear (U. arctos) uncovered offset signals in timing of population expansion between the two species, that are attributed to differential demographic responses to past climate cycling. Mitogenomic structure of polar bears was shallow and developed recently, in contrast to the multiple clades of brown bears. We found no genetic signatures of recent hybridization between the species in our large, circumpolar sample, suggesting that recently observed hybrids represent localized events. Documenting changes in subpopulation connectivity will allow polar nations to proactively adjust conservation actions to continuing decline in sea-ice habitat.     

Estimating the Barents Sea polar bear subpopulation size

A large-scale survey was conducted in August 2004 to estimate the size of the Barents Sea polar bear subpopulation. We combined helicopter line transect distance sampling (DS) surveys in most of the survey area with total counts in small areas not suitable for DS. Due to weather constraints we failed to survey some of the areas originally planned to be covered by DS. For those, abundance was estimated using a ratio estimator, in which the auxiliary variable was the number of satellite telemetry fixes (in previous years). We estimated that the Barents Sea subpopulation had approximately 2,650 (95% CI approximately 1,900–3,600) bears. Given current intense interest in polar bear management due to the potentially disastrous effects of climate change, it is surprising that many subpopulation sizes are still unknown. We show here that line transect sampling is a promising method for addressing the need for abundance estimates.     

Eisbärenzucht (Ursus maritimus Phipps, 1774) im Zoo am Meer Bremerhaven – Fortsetzung einer langen Tradition

Breeding polar bears has a long tradition in the Zoo am Meer, the “Zoo at the Sea” in Bremerhaven. In the years between 1935 and 1974 three female bears gave birth to 32 cubs and 19 of them were successfully raised. In the following years many attempts to tie in with this tradition failed. Bad constructional conditions and disadvantageous animal pairings were the main reasons for the missing breeding success. After completely reconstructing the polar bear enclosure in the year 2002, as well as extending the breeding area with an additional smaller breeding cave in 2012 and pairing male Lloyd and female Valeska in 2013 a new polar bear birth and upbringing could be experienced after 40 years. Offspring Lale is the 35^th cub in Bremerhaven Zoo.     

Cranial distinctiveness in the Apennine brown bear: genetic drift effect or ecophenotypic adaptation?

Molecular studies highlighted a strong genetic affinity between the remnant and isolated population of the Apennine brown bear and other southern European populations. Despite this genetic closeness a recent morphometric study revealed a marked phenotypic distinctiveness of the Apennine population, supporting the reinstatement of a distinct taxon (Ursus arctos marsicanus). By building upon previous morphological analyses, we adopted geometric morphometrics to better investigate the skull morphology of the Apennine brown bear with reference to the other, closely related southern European populations. Both skull shape and size differences confirmed the strong divergence of U. arctos marsicanus. In particular, the Apennine bears are characterized by an enlargement of the supraorbital apophysis and a larger distance across the zygomatic arches. Furthermore, our analyses highlighted significant shape differences of the first upper molar in the Apennine bears. Our results suggest that the Apennine bears underwent a rapid morphological change, possibly driven by genetic drift and local selective pressures. Because the greatest morphological differentiation is likely to be related to the muscles involved in mastication, we hypothesize that local selective pressures might be related to a shift in food habits, with highly reduced depredation and feeding on large carcasses in favour of vegetation and hard mast (beech nuts and acorns). These results suggest an adaptive distinctiveness of the Apennine bears, which should be carefully considered in any management and conservation action addressed to this highly endangered population. Although more in‐depth molecular studies are required to better assess the taxonomic and genetic status of the Apennine brown bear population, our study emphasizes the importance of morphological analyses as a complementary tool for a more thorough characterization of variation and divergence in endangered taxa. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Recent observations of intraspecific predation and cannibalism among polar bears in the southern Beaufort Sea

Intraspecific killing has been reported among polar bears (Ursus maritimus), brown bears (U. arctos), and black bears (U. americanus). Although cannibalism is one motivation for such killings, the ecological factors mediating such events are poorly understood. Between 24 January and 10 April 2004, we confirmed three instances of intraspecific predation and cannibalism in the Beaufort Sea. One of these, the first of this type ever reported for polar bears, was a parturient female killed at her maternal den. The predating bear was hunting in a known maternal denning area and apparently discovered the den by scent. A second predation event involved an adult female and cub recently emerged from their den, and the third involved a yearling male. During 24 years of research on polar bears in the southern Beaufort Sea region of northern Alaska and 34 years in northwestern Canada, we have not seen other incidents of polar bears stalking, killing, and eating other polar bears. We hypothesize that nutritional stresses related to the longer ice-free seasons that have occurred in the Beaufort Sea in recent years may have led to the cannibalism incidents we observed in 2004.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Red king crab (<Emphasis Type="Italic">Paralithodes camtschaticus</Emphasis>) in the Barents Sea: A comparative study of introduced and native populations

Red king crab (Paralithodes camtschaticus) was introduced into the Barents Sea in the 1960–1970s. At present, it occurs along the coast from Hammerfest (Northern Norway) to the (northeastern coast of Kola Peninsula). We studied the polymorphism of a mitochondrial gene encoding cytochrome oxidase (COI) and five nuclear microsatellite loci in four samples from the Barents Sea and two donor populations from the Western Kamchatka and Primorye (Russian Pacifics). No decrease in the genetic diversity of the introduced populations was observed. The results of PCA103 locusanalysis but some samples from the Barents Sea significantly differed from the Pacific populations according to the test for population differentiation demonstrated that the sample from Varrangerfjord was highly significantly different from the other five populations. As no significant differences between the other samples were found at this locus. We consider it to be a marker for Varangerfjord population differentiation. Possible reasons of this phenomenon are discussed.     

Population substructure and space use of Foxe Basin polar bears

Climate change has been identified as a major driver of habitat change, particularly for sea ice-dependent species such as the polar bear (Ursus maritimus). Population structure and space use of polar bears have been challenging to quantify because of their circumpolar distribution and tendency to range over large areas. Knowledge of movement patterns, home range, and habitat is needed for conservation and management. This is the first study to examine the spatial ecology of polar bears in the Foxe Basin management unit of Nunavut, Canada. Foxe Basin is in the mid-Arctic, part of the seasonal sea ice ecoregion and it is being negatively affected by climate change. Our objectives were to examine intrapopulation spatial structure, to determine movement patterns, and to consider how polar bear movements may respond to changing sea ice habitat conditions. Hierarchical and fuzzy cluster analyses were used to assess intrapopulation spatial structure of geographic position system satellite-collared female polar bears. Seasonal and annual movement metrics (home range, movement rates, time on ice) and home-range fidelity (static and dynamic overlap) were compared to examine the influence of regional sea ice on movements. The polar bears were distributed in three spatial clusters, and there were differences in the movement metrics between clusters that may reflect sea ice habitat conditions. Within the clusters, bears moved independently of each other. Annual and seasonal home-range fidelity was observed, and the bears used two movement patterns: on-ice range residency and annual migration. We predict that home-range fidelity may decline as the spatial and temporal predictability of sea ice changes. These new findings also provide baseline information for managing and monitoring this polar bear population.     

Geographic variation of PCB congeners in polar bears (Ursus maritimus) from Svalbard east to the Chukchi Sea

We present data on geographic variation in polychlorinated biphenyl (PCB) congeners in adult female polar bears (Ursus maritimus) from Svalbard eastward to the Chukchi Sea. Blood samples from 90 free-living polar bears were collected in 1987-1995. Six PCB congeners, penta to octa chlorinated (PCB-99, -118, -153, -156, -180, -194), were selected for this study. Differences between areas were found in PCB levels and congener patterns. Bears from Franz Josef Land (11,194 ng/g lipid weight) and the Kara Sea (9,412 ng/g lw) had similar DPCB levels and were higher than all other populations (Svalbard 5,043 ng/g lw, East Siberian Sea 3,564 ng/g lw, Chukchi Sea 2,465 ng/g lw). Svalbard PCB levels were higher than those from the Chukchi Sea. Our results, combined with earlier findings, indicate that polar bears from Franz Josef Land and the Kara Sea have the highest PCB levels in the Arctic. Decreasing trends were seen eastwards and westwards from this region. Of the congeners investigated in the present study, the lower chlorinated PCBs are increasing and the high chlorinated PCBs are decreasing from Svalbard eastward to the Chukchi Sea. Different pollution sources, compound transport patterns and regional prey differences could explain the variation in PCB congener levels and patterns between regions.     

Genetic status of Asiatic black bear (Ursus thibetanus) reintroduced into South Korea based on mitochondrial DNA and microsatellite loci analysis

The Asiatic black bear is one of the most endangered mammals in South Korea owing to population declines resulting from human exploitation and habitat fragmentation. To restore the black bear population in South Korea, 27 bear cubs from North Korea and Russian Far East (Primorsky Krai) were imported and released into Jirisan National Park, a reservoir of the largest wild population in South Korea, in 2004. To monitor the success of this reintroduction, the genetic diversity and population structure of the reintroduced black bears were measured using both mitochondrial and nuclear DNA markers. Mitochondrial D-loop region DNA sequences (615 bp) of 43 Japanese black bears from previous study and 14 Southeast Asian black bears in this study were employed to obtain phylogenetic inference of the reintroduced black bears. The mitochondrial phylogeny indicated Asiatic black bear populations from Russian Far East and North Korea form a single evolutionary unit distinct from populations from Japan and Southeast Asia. Mean expected heterozygosity (H(E)) across 16 microsatellite loci was 0.648 for Russian and 0.676 for North Korean populations. There was a moderate but significant level of microsatellite differentiation (F(ST) = 0.063) between black bears from the 2 source areas. In addition, genetic evidences revealed that 2 populations are represented as diverging groups, with lingering genetic admixture among individuals of 2 source populations. Relatedness analysis based on genetic markers indicated several discrepancies with the pedigree records. Implication of the phylogenetic and genetic evidences on long-term management of Asiatic black bears in South Korea is discussed.