Determining causes of genetic isolation in a large carnivore (Ursus americanus) population to direct contemporary conservation measures.

The processes leading to genetic isolation influence a population's local extinction risk, and should thus be identified before conservation actions are implemented. Natural or human-induced circumstances can result in historical or contemporary barriers to gene flow and/or demographic bottlenecks. Distinguishing between these hypotheses can be achieved by comparing genetic diversity and differentiation in isolated vs. continuous neighboring populations. In Ontario, American black bears (Ursus americanus) are continuously distributed, genetically diverse, and exhibit an isolation-by-distance structuring pattern, except on the Bruce Peninsula (BP). To identify the processes that led to the genetic isolation of BP black bears, we modelled various levels of historical and contemporary migration and population size reductions using forward simulations. We compared simulation results with empirical genetic indices from Ontario black bear populations under different levels of geographic isolation, and conducted additional simulations to determine if translocations could help achieve genetic restoration. From a genetic standpoint, conservation concerns for BP black bears are warranted because our results show that: i) a recent demographic bottleneck associated with recently reduced migration best explains the low genetic diversity on the BP; and ii) under sustained isolation, BP black bears could lose between 70% and 80% of their rare alleles within 100 years. Although restoring migration corridors would be the most effective method to enhance long-term genetic diversity and prevent inbreeding, it is unrealistic to expect connectivity to be re-established. Current levels of genetic diversity could be maintained by successfully translocating 10 bears onto the peninsula every 5 years. Such regular translocations may be more practical than landscape restoration, because areas connecting the peninsula to nearby mainland black bear populations have been irreversibly modified by humans, and form strong barriers to movement.

Comparison of methanol and isopropanol as wash solvents for determination of hair cortisol concentration in grizzly bears and polar bears.

Methodological differences among laboratories are recognized as significant sources of variation in quantification of hair cortisol concentration (HCC). An important step in processing hair, particularly when collected from wildlife, is the choice of solvent used to remove or "wash" external hair shaft cortisol prior to quantification of HCC. The present study systematically compared methanol and isopropanol as wash solvents for their efficiency at removing external cortisol without extracting internal hair shaft cortisol in samples collected from free-ranging grizzly bears and polar bears. Cortisol concentrations in solvents and hair were determined in each of one to eight washes of hair with each solvent independently. •There were no significant decreases in internal hair shaft cortisol among all eight washes for either solvent, although methanol removed detectable hair surface cortisol after one wash in grizzly bear hair whereas hair surface cortisol was detected in all eight isopropanol washes.•There were no significant differences in polar bear HCC washed one to eight times with either solvent, but grizzly bear HCC was significantly greater in hair washed with isopropanol compared to methanol.•There were significant differences in HCC quantified using different commercial ELISA kits commonly used for HCC determinations.

Assessing polar bear (Ursus maritimus) population structure in the Hudson Bay region using SNPs.

Defining subpopulations using genetics has traditionally used data from microsatellite markers to investigate population structure; however, single-nucleotide polymorphisms (SNPs) have emerged as a tool for detection of fine-scale structure. In Hudson Bay, Canada, three polar bear (Ursus maritimus) subpopulations (Foxe Basin (FB), Southern Hudson Bay (SH), and Western Hudson Bay (WH)) have been delineated based on mark-recapture studies, radiotelemetry and satellite telemetry, return of marked animals in the subsistence harvest, and population genetics using microsatellites. We used SNPs to detect fine-scale population structure in polar bears from the Hudson Bay region and compared our results to the current designations using 414 individuals genotyped at 2,603 SNPs. Analyses based on discriminant analysis of principal components (DAPC) and STRUCTURE support the presence of four genetic clusters: (i) Western-including individuals sampled in WH, SH (excluding Akimiski Island in James Bay), and southern FB (south of Southampton Island); (ii) Northern-individuals sampled in northern FB (Baffin Island) and Davis Strait (DS) (Labrador coast); (iii) Southeast-individuals from SH (Akimiski Island in James Bay); and (iv) Northeast-individuals from DS (Baffin Island). Population structure differed from microsatellite studies and current management designations demonstrating the value of using SNPs for fine-scale population delineation in polar bears.

Conservation status of polar bears (Ursus maritimus) in relation to projected sea-ice declines.

Loss of Arctic sea ice owing to climate change is the primary threat to polar bears throughout their range. We evaluated the potential response of polar bears to sea-ice declines by (i) calculating generation length (GL) for the species, which determines the timeframe for conservation assessments; (ii) developing a standardized sea-ice metric representing important habitat; and (iii) using statistical models and computer simulation to project changes in the global population under three approaches relating polar bear abundance to sea ice. Mean GL was 11.5 years. Ice-covered days declined in all subpopulation areas during 1979-2014 (median -1.26 days year-1). The estimated probabilities that reductions in the mean global population size of polar bears will be greater than 30%, 50% and 80% over three generations (35-41 years) were 0.71 (range 0.20-0.95), 0.07 (range 0-0.35) and less than 0.01 (range 0-0.02), respectively. According to IUCN Red List reduction thresholds, which provide a common measure of extinction risk across taxa, these results are consistent with listing the species as vulnerable. Our findings support the potential for large declines in polar bear numbers owing to sea-ice loss, and highlight near-term uncertainty in statistical projections as well as the sensitivity of projections to different plausible assumptions.

Does Predation Influence the Seasonal and Diel Timing of Moose Calving in Central Ontario, Canada?

Birth synchrony is well documented among ungulates and is hypothesised to maximize neonate survival, either by minimizing the risk of predation through predator swamping or by synchronising birthing with increased seasonal food availability. We used encapsulated vaginal implant transmitters to locate and capture neonatal moose calves and document the seasonal and diel timing of parturition in two adjacent study areas with different predation pressure in central Ontario, Canada. We tested the hypothesis that predation promotes earlier and more synchronous birth of moose calves. Across both areas, proportionately more births occurred during the afternoon and fewer than expected occurred overnight. Mean date of calving averaged 1.5 days earlier and calving was also more synchronous in the study area with heavier predation pressure, despite average green-up date and peak Normalized Difference Vegetation Index date occurring 2 days later in this study area than in the area receiving lighter predation pressure. We encourage analysis of data on timing of parturition from additional study areas experiencing varying degrees of predation pressure to better clarify the influence of predation in driving seasonal and diel timing of parturition in temperate ungulates.

Establishing a definition of polar bear (Ursus maritimus) health: a guide to research and management activities.

The meaning of health for wildlife and perspectives on how to assess and measure health, are not well characterized. For wildlife at risk, such as some polar bear (Ursus maritimus) subpopulations, establishing comprehensive monitoring programs that include health status is an emerging need. Environmental changes, especially loss of sea ice habitat, have raised concern about polar bear health. Effective and consistent monitoring of polar bear health requires an unambiguous definition of health. We used the Delphi method of soliciting and interpreting expert knowledge to propose a working definition of polar bear health and to identify current concerns regarding health, challenges in measuring health, and important metrics for monitoring health. The expert opinion elicited through the exercise agreed that polar bear health is defined by characteristics and knowledge at the individual, population, and ecosystem level. The most important threats identified were in decreasing order: climate change, increased nutritional stress, chronic physiological stress, harvest management, increased exposure to contaminants, increased frequency of human interaction, diseases and parasites, and increased exposure to competitors. Fifteen metrics were identified to monitor polar bear health. Of these, indicators of body condition, disease and parasite exposure, contaminant exposure, and reproductive success were ranked as most important. We suggest that a cumulative effects approach to research and monitoring will improve the ability to assess the biological, ecological, and social determinants of polar bear health and provide measurable objectives for conservation goals and priorities and to evaluate progress.

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.

Polar and brown bear genomes reveal ancient admixture and demographic footprints of past climate change.

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.

Incidental nest predation in freshwater turtles: inter- and intraspecific differences in vulnerability are explained by relative crypsis.

There has long been interest in the influence of predators on prey populations, although most predator-prey studies have focused on prey species that are targets of directed predator searching. Conversely, few have addressed depredation that occurs after incidental encounters with predators. We tested two predictions stemming from the hypothesis that nest predation on two sympatric freshwater turtle species whose nests are differentially prone to opportunistic detection-painted turtles (Chrysemys picta) and snapping turtles (Chelydra serpentina)-is incidental: (1) predation rates should be density independent, and (2) individual predators should not alter their foraging behavior after encountering nests. After monitoring nest survival and predator behavior following nest depredation over 2 years, we confirmed that predation by raccoons (Procyon lotor), the primary nest predators in our study area, matched both predictions. Furthermore, cryptic C. picta nests were victimized with lower frequency than more detectable C. serpentina nests, and nests of both species were more vulnerable in human-modified areas where opportunistic nest discovery is facilitated. Despite apparently being incidental, predation on nests of both species was intensive (57% for painted turtles, 84% for snapping turtles), and most depredations occurred within 1 day of nest establishment. By implication, predation need not be directed to affect prey demography, and factors influencing prey crypsis are drivers of the impact of incidental predation on prey. Our results also imply that efforts to conserve imperiled turtle populations in human-modified landscapes should include restoration of undisturbed conditions that are less likely to expose nests to incidental predators.

Serum corticosteroid binding globulin expression is modulated by fasting in polar bears (Ursus maritimus).

Polar bears (Ursus maritimus) from several subpopulations undergo extended fasting during the ice-free season. However, the animals appear to conserve protein despite the prolonged fasting, though the mechanisms involved are poorly understood. We hypothesized that elevated concentrations of corticosteroid binding globulin (CBG), the primary cortisol binding protein in circulation, lead to cortisol resistance and provide a mechanism for protein conservation during extended fasting. The metabolic state (feeding vs. fasting) of 16 field sampled male polar bears was determined based on their serum urea to creatinine ratio (>25 for feeding vs. <5 for fasting). There were no significant differences in serum cortisol levels between all male and female polar bears sampled. Serum CBG expression was greater in lactating females relative to non-lactating females and males. CBG expression was significantly higher in fasting males when compared to non-fasting males. This leads us to suggest that CBG expression may serve as a mechanism to conserve protein during extended fasting in polar bears by reducing systemic free cortisol concentrations. This was further supported by a lower serum glucose concentration in the fasting bears. As well, a lack of an enhanced adrenocortical response to acute capture stress supports our hypothesis that chronic hunger is not a stressor in this species. Overall, our results suggest that elevated serum CBG expression may be an important adaptation to spare proteins by limiting cortisol bioavailability during extended fasting in polar bears.

Use of hyaluronidase to improve chemical immobilization of free-ranging polar bears (Ursus maritimus).

We assessed the efficacy and safety of hyaluronidase to improve chemical immobilization of free-ranging polar bears (Ursus maritimus) captured from helicopter by remote drug delivery along the Ontario coast line of northwestern James Bay and southern Hudson Bay during September 2005 and October 2007. We used a single blind study design in which one person prepared and loaded all darts without the shooter knowing whether hyaluronidase (150 IU per dart) or sterile water was added to the immobilizing drug mixture of xylazine and zolazepam-tiletamine (XZT). We found that we often required more than one dart to immobilize bears in the control group (XZT+sterile water; >1 dart for 15 of 28 captures) versus the treatment group (XZT+hyaluronidase; >1 dart for seven of 26 captures). As a consequence, treatment bears were generally immobilized with smaller XZT dosages (7.9 vs. 9.4 mg/kg; P = 0.08) and shorter induction (10 vs. 15 min; P = 0.004) than control bears. We found no differences in vital rates and serum biochemistry results between control and treatment bears. We did find, however, that induction times correlated directly with rectal temperature at

A body composition model to estimate mammalian energy stores and metabolic rates from body mass and body length, with application to polar bears.

Many 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, non-invasive and provides considerably more information on the energetic status of individuals than currently available methods.