Polar bears (PBs) are superbly adapted to the extreme Arctic environment and have become emblematic of the threat to biodiversity from global climate change. Their divergence from the lower-latitude brown bear provides a textbook example of rapid evolution of distinct phenotypes. However, limited mitochondrial and nuclear DNA evidence conflicts in the timing of PB origin as well as placement of the species within versus sister to the brown bear lineage. We gathered extensive genomic sequence data from contemporary polar, brown, and American black bear samples, in addition to a 130,000-to 110,000-y old PB, to examine this problem from a genome-wide perspective. Nuclear DNA markers reflect a species tree consistent with expectation, showing polar and brown bears to be sister species. However, for the enigmatic brown bears native to Alaska's Alexander Archipelago, we estimate that not only their mitochondrial genome, but also 5-10% of their nuclear genome, is most closely related to PBs, indicating ancient admixture between the two species. Explicit admixture analyses are consistent with ancient splits among PBs, brown bears and black bears that were later followed by occasional admixture. We also provide paleodemographic estimates that suggest bear evolution has tracked key climate events, and that PB in particular experienced a prolonged and dramatic decline in its effective population size during the last ca. 500,000 years. We demonstrate that brown bears and PBs have had sufficiently independent evolutionary histories over the last 4-5 million years to leave imprints in the PB nuclear genome that likely are associated with ecological adaptation to the Arctic environment.demographic history | hybridization | mammalian genomics | phylogenetics G enome-scale studies of speciation and admixture have become essential tools in evolutionary analyses of recently diverged lineages. For example, paradigm-shifting genomic research on archaic and anatomically modern humans has identified critical gene flow events during hominin history (1, 2). However, aside from several analyses of domesticated species and their wild relatives (e.g., ref.3), studies that use whole-genome sequencing to investigate admixture in wildlife populations are only now beginning to emerge.The bear family (Ursidae, Mammalia) represents an excellent, largely untapped model for investigating complex speciation and rapid evolution of distinct phenotypes. Although polar bears (PBs; Ursus maritimus) and brown bears (Ursus arctos) are considered separate species, analyses of fossil evidence and mitochondrial sequence data have indicated a recent divergence of PBs from within brown bears (surveyed in ref. 4). For example, phylogenetic analyses of complete mitochondrial genomes, including from a unique 130,000-to 110,000-y-old PB jawbone from Svalbard, Norway, confirmed a particularly close relationship between PB and a genetically isolated population of brown bears from the Admiralty, Baranof, and Chichagof islands in Alaska's Alexander Archipelago (hereaf...

► Unpublished and published data were compiled for Arctic fish, birds, and mammals. ► These data were compared to available toxicological threshold limits. ► Toothed whales, polar bears, and some bird and fish species exceeded the limits. ► Increasing mercury concentrations are observed for some Arctic species. ► These exceeded thresholds and increasing Hg trends are of concern. a b s t r a c t a r t i c l e i n f o This review critically evaluates the available mercury (Hg) data in Arctic marine biota and the Inuit population against toxicity threshold values. In particular marine top predators exhibit concentrations of mercury in their tissues and organs that are believed to exceed thresholds for biological effects. Species whose concentrations exceed threshold values include the polar bears (Ursus maritimus), beluga whale (Delphinapterus leucas), pilot whale (Globicephala melas), hooded seal (Cystophora cristata), a few seabird species, and landlocked Arctic char (Salvelinus alpinus). Toothed whales appear to be one of the most vulnerable groups, with high concentrations Science of the Total Environment 443 (2013) [775][776][777][778][779][780][781][782][783][784][785][786][787][788][789][790]

Summary1. Population density is a critical ecological parameter informing effective wildlife management and conservation decisions. Density is often estimated by dividing capture-recapture (C-R) estimates of abundance (N) by size of the study area, but this relies on the assumption of geographic closure -a situation rarely achieved in studies of large carnivores. For geographically open populationsN is overestimated relative to the size of the study area because animals with only part of their home range on the study area are available for capture. This bias ('edge effect') is more severe when animals such as large carnivores range widely. To compensate for edge effect, a boundary strip around the trap array is commonly included when estimating the effective trap area (Â). Various methods for estimating the width of the boundary strip are proposed, butN ⁄ estimates of large carnivore density are generally mistrusted unless concurrent telemetry data are available to defineÂ. Remote sampling by cameras or hair snags may reduce study costs and duration, yet without telemetry data inflated density estimates remain problematic. 2. We evaluated recently developed spatially explicit capture-recapture (SECR) models using data from a common large carnivore, the American black bear Ursus americanus, obtained by remote sampling of 11 geographically open populations. These models permit direct estimation of population density from C-R data without assuming geographic closure. We compared estimates derived using this approach to those derived using conventional approaches that estimate density asN ⁄Â. 3. Spatially explicit C-R estimates were 20-200% lower than densities estimated asN ⁄Â. AIC c supported individual heterogeneity in capture probabilities and home range sizes. Variable home range size could not be accounted for when estimating density asN ⁄Â. 4. Synthesis and applications. We conclude that the higher densities estimated asN ⁄ compared to estimates from SECR models are consistent with positive bias due to edge effects in the former. Inflated density estimates could lead to management decisions placing threatened or endangered large carnivores at greater risk. Such decisions could be avoided by estimating density by SECR when bias due to geographic closure violation cannot be minimized by study design.

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.

SUMMARYMany species experience large fluctuations in food availability and depend on energy from fat and protein stores for survival, reproduction and growth. Body condition and, more specifically, energy stores thus constitute key variables in the life history of many species. Several indices exist to quantify body condition but none can provide the amount of stored energy. To estimate energy stores in mammals, we propose a body composition model that differentiates between structure and storage of an animal. We develop and parameterize the model specifically for polar bears (Ursus maritimus Phipps) but all concepts are general and the model could be easily adapted to other mammals. The model provides predictive equations to estimate structural mass, storage mass and storage energy from an appropriately chosen measure of body length and total body mass. The model also provides a means to estimate basal metabolic rates from body length and consecutive measurements of total body mass. Model estimates of body composition, structural mass, storage mass and energy density of 970 polar bears from Hudson Bay were consistent with the life history and physiology of polar bears. Metabolic rate estimates of fasting adult males derived from the body composition model corresponded closely to theoretically expected and experimentally measured metabolic rates. Our method is simple, noninvasive and provides considerably more information on the energetic status of individuals than currently available methods.

In this investigation a body-condition index (BCI) was developed for polar bears (Ursus maritimus), black bears (Ursus americanus), and grizzly bears (Ursus arctos), based on residuals from the regression of total body mass against a linear measure of size, straight-line body length (SLBL). Transformation of mass–length data from 1198 polar bears, 595 black bears, and 126 grizzly bears to natural logarithms resulted in a linear relationship between mass and length. However, the relationship in polar bears differed from that in black and grizzly bears. SLBL had a close positive relationship with skeletal (bone) mass in polar bears (n = 31) and black bears (n = 33), validating the use of SLBL as an accurate index of body size. There was no correlation between SLBL and BCI for polar bears (r = 0.005, p = 0.87, n = 1198) or for black bears and grizzly bears (r = 0.04, p = 0.30, n = 721), indicating that the BCI was independent of body size. The BCI had a close positive relationship with true body condition, measured as the standardized residual of the combined mass of fat and skeletal muscle against SLBL, in polar and black bears that were dissected to determine individual tissue masses. The BCI also had a close positive relationship with the standardized residual of fat mass against SLBL. Estimation of BCI values for polar bears, or for black bears and grizzly bears, is facilitated by prediction equations that require measurement of total body mass and SLBL for individual animals.

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Long-term physiological stress in individual animals may be an important mechanism linking ecological change with impaired wildlife population health. In the Southern Hudson Bay (SH) subpopulation of polar bears (Ursus maritimus), increasing stress associated with climate warming may be related to declining body condition. Accordingly, the development of tools to assess long-term stress in this species may prove invaluable for conservation efforts in this threatened population. The measurement of hair cortisol concentration (HCC) has shown promise as a potential biomarker of long-term stress in free-ranging bears. However, to serve as a useful management tool, factors influencing HCC in polar bears must be identified and then revealed to establish linkages between environmental conditions and the fitness of individual animals. We determined HCC (median ¼ 0.48 pg/mg [range ¼ 0.16-2.26 pg/mg]) in 185 polar bears captured in southern Hudson Bay from 2007 to 2009. HCC was influenced by sex, family group status, and capture period but not by body region or hair type. Using models developed through a combination of hypothesis testing and information theory, we also determined that HCC was negatively associated with growth indices (length, mass, and body condition index) linked to fitness in polar bears. Additional research will be required across several polar bear populations to establish the utility of HCC as a tool for polar bear conservation. ß 2012 The Wildlife Society.