A polar bear paleogenome reveals extensive ancient gene flow from polar bears into brown bears.

Polar bears (Ursus maritimus) and brown bears (Ursus arctos) are sister species possessing distinct physiological and behavioural adaptations that evolved over the last 500,000 years. However, comparative and population genomics analyses have revealed that several extant and extinct brown bear populations have relatively recent polar bear ancestry, probably as the result of geographically localized instances of gene flow from polar bears into brown bears. Here, we generate and analyse an approximate 20X paleogenome from an approximately 100,000-year-old polar bear that reveals a massive prehistoric admixture event, which is evident in the genomes of all living brown bears. This ancient admixture event was not visible from genomic data derived from living polar bears. Like more recent events, this massive admixture event mainly involved unidirectional gene flow from polar bears into brown bears and occurred as climate changes caused overlap in the ranges of the two species. These findings highlight the complex reticulate paths that evolution can take within a regime of radically shifting climate.

Influence of sea ice dynamics on population energetics of Western Hudson Bay polar bears.

The Arctic marine ecosystem has experienced extensive changes in sea ice dynamics, with significant effects on ice-dependent species such as polar bears (Ursus maritimus). We used annual estimates of the numbers of bears onshore in the core summering area, age/sex structure and body condition data to estimate population energy density and storage energy in Western Hudson Bay polar bears from 1985 to 2018. We examined intra-population variation in energetic patterns, temporal energetic trends and the relationship between population energetics and sea ice conditions. Energy metrics for most demographic classes declined over time in relation to earlier sea ice breakup, most significantly for solitary adult females and subadult males, suggesting their greater vulnerability to nutritional stress than other age/sex classes. Temporal declines in population energy metrics were related to earlier breakup and longer lagged open-water periods, suggesting multi-year effects of sea ice decline. The length of the open-water period ranged from 102 to 166 days and increased significantly by 9.9 days/decade over the study period. Total population energy density and storage energy were significantly lower when sea ice breakup occurred earlier and the lagged open-water period was longer. At the earliest breakup and a lagged open-water period of 180 days, population energy density was predicted to be 33% lower than our minimum estimated energy density and population storage energy was predicted to be 40% lower than the minimum estimated storage energy. Consequently, over the study, the total population energy density declined by 53% (mean: 3668 ± 386 MJ kg-1/decade) and total population storage energy declined by 56% (mean: 435900 ± 46770 MJ/decade). This study provides insights into ecological mechanisms linking population responses to sea ice decline and highlights the significance of maintaining long-term research programs.

Corrigendum: Internet Blogs, Polar Bears, and Climate-Change Denial by Proxy.

[This corrects the article DOI: 10.1093/biosci/bix133.].

Internet Blogs, Polar Bears, and Climate-Change Denial by Proxy.

Increasing surface temperatures, Arctic sea-ice loss, and other evidence of anthropogenic global warming (AGW) are acknowledged by every major scientific organization in the world. However, there is a wide gap between this broad scientific consensus and public opinion. Internet blogs have strongly contributed to this consensus gap by fomenting misunderstandings of AGW causes and consequences. Polar bears (Ursus maritimus) have become a "poster species" for AGW, making them a target of those denying AGW evidence. Here, focusing on Arctic sea ice and polar bears, we show that blogs that deny or downplay AGW disregard the overwhelming scientific evidence of Arctic sea-ice loss and polar bear vulnerability. By denying the impacts of AGW on polar bears, bloggers aim to cast doubt on other established ecological consequences of AGW, aggravating the consensus gap. To counter misinformation and reduce this gap, scientists should directly engage the public in the media and blogosphere.

Genomic Evidence of Widespread Admixture from Polar Bears into Brown Bears during the Last Ice Age.

Recent genomic analyses have provided substantial evidence for past periods of gene flow from polar bears (Ursus maritimus) into Alaskan brown bears (Ursus arctos), with some analyses suggesting a link between climate change and genomic introgression. However, because it has mainly been possible to sample bears from the present day, the timing, frequency, and evolutionary significance of this admixture remains unknown. Here, we analyze genomic DNA from three additional and geographically distinct brown bear populations, including two that lived temporally close to the peak of the last ice age. We find evidence of admixture in all three populations, suggesting that admixture between these species has been common in their recent evolutionary history. In addition, analyses of ten fossil bears from the now-extinct Irish population indicate that admixture peaked during the last ice age, whereas brown bear and polar bear ranges overlapped. Following this peak, the proportion of polar bear ancestry in Irish brown bears declined rapidly until their extinction. Our results support a model in which ice age climate change created geographically widespread conditions conducive to admixture between polar bears and brown bears, as is again occurring today. We postulate that this model will be informative for many admixing species pairs impacted by climate change. Our results highlight the power of paleogenomics to reveal patterns of evolutionary change that are otherwise masked in contemporary data.

Spring fasting behavior in a marine apex predator provides an index of ecosystem productivity.

The effects of declining Arctic sea ice on local ecosystem productivity are not well understood but have been shown to vary inter-specifically, spatially, and temporally. Because marine mammals occupy upper trophic levels in Arctic food webs, they may be useful indicators for understanding variation in ecosystem productivity. Polar bears (Ursus maritimus) are apex predators that primarily consume benthic and pelagic-feeding ice-associated seals. As such, their productivity integrates sea ice conditions and the ecosystem supporting them. Declining sea ice availability has been linked to negative population effects for polar bears but does not fully explain observed population changes. We examined relationships between spring foraging success of polar bears and sea ice conditions, prey productivity, and general patterns of ecosystem productivity in the Beaufort and Chukchi Seas (CSs). Fasting status (≥7 days) was estimated using serum urea and creatinine levels of 1,448 samples collected from 1,177 adult and subadult bears across three subpopulations. Fasting increased in the Beaufort Sea between 1983-1999 and 2000-2016 and was related to an index of ringed seal body condition. This change was concurrent with declines in body condition of polar bears and observed changes in the diet, condition and/or reproduction of four other vertebrate consumers within the food chain. In contrast, fasting declined in CS polar bears between periods and was less common than in the two Beaufort Sea subpopulations consistent with studies demonstrating higher primary productivity and maintenance or improved body condition in polar bears, ringed seals, and bearded seals despite recent sea ice loss in this region. Consistency between regional and temporal variation in spring polar bear fasting and food web productivity suggests that polar bears may be a useful indicator species. Furthermore, our results suggest that spatial and temporal ecological variation is important in affecting upper trophic-level productivity in these marine ecosystems.

Demography of an apex predator at the edge of its range: impacts of changing sea ice on polar bears in Hudson Bay.

Changes in the abundance and distribution of wildlife populations are common consequences of historic and contemporary climate change. Some Arctic marine mammals, such as the polar bear (Ursus maritimus), may be particularly vulnerable to such changes due to the loss of Arctic sea ice. We evaluated the impacts of environmental variation on demographic rates for the Western Hudson Bay (WH), polar bear subpopulation from 1984 to 2011 using live-recapture and dead-recovery data in a Bayesian implementation of multistate capture-recapture models. We found that survival of female polar bears was related to the annual timing of sea ice break-up and formation. Using estimated vital rates (e.g., survival and reproduction) in matrix projection models, we calculated the growth rate of the WH subpopulation and projected population responses under different environmental scenarios while accounting for parametric uncertainty, temporal variation, and demographic stochasticity. Our analysis suggested a long-term decline in the number of bears from 1185 (95% Bayesian credible interval [BCI] = 993-1411) in 1987 to 806 (95% BCI = 653-984) in 2011. In the last 10 yr of the study, the number of bears appeared stable due to temporary stability in sea ice conditions (mean population growth rate for the period 2001-2010 = 1.02, 95% BCI = 0.98-1.06). Looking forward, we estimated long-term growth rates for the WH subpopulation of ~1.02 (95% BCI = 1.00-1.05) and 0.97 (95% BCI = 0.92-1.01) under hypothetical high and low sea ice conditions, respectively. Our findings support previous evidence for a demographic linkage between sea ice conditions and polar bear population dynamics. Furthermore, we present a robust framework for sensitivity analysis with respect to continued climate change (e.g., to inform scenario planning) and for evaluating the combined effects of climate change and management actions on the status of wildlife populations.

Mass Loss Rates of Fasting Polar Bears.

Polar bears (Ursus maritimus) have adapted to an annual cyclic regime of feeding and fasting, which is extreme in seasonal sea ice regions of the Arctic. As a consequence of climate change, sea ice breakup has become earlier and the duration of the open-water period through which polar bears must rely on fat reserves has increased. To date, there is limited empirical data with which to evaluate the potential energetic capacity of polar bears to withstand longer fasts. We measured the incoming and outgoing mass of inactive polar bears (n = 142) that were temporarily detained by Manitoba Conservation and Water Stewardship during the open-water period near the town of Churchill, Manitoba, Canada, in 2009-2014. Polar bears were given access to water but not food and held for a median length of 17 d. Median mass loss rates were 1.0 kg/d, while median mass-specific loss rates were 0.5%/d, similar to other species with high adiposity and prolonged fasting capacities. Mass loss by unfed captive adult males was identical to that lost by free-ranging individuals, suggesting that terrestrial feeding contributes little to offset mass loss. The inferred metabolic rate was comparable to a basal mammalian rate, suggesting that while on land, polar bears can maintain a depressed metabolic rate to conserve energy. Finally, we estimated time to starvation for subadults and adult males for the on-land period. Results suggest that at 180 d of fasting, 56%-63% of subadults and 18%-24% of adult males in this study would die of starvation. Results corroborate previous assessments on the limits of polar bear capacity to withstand lengthening ice-free seasons and emphasize the greater sensitivity of subadults to changes in sea ice phenology.

Polar bear population dynamics in the southern Beaufort Sea during a period of sea ice decline.

In the southern Beaufort Sea of the United States and Canada, prior investigations have linked declines in summer sea ice to reduced physical condition, growth, and survival of polar bears (Ursus maritimus). Combined with projections of population decline due to continued climate warming and the ensuing loss of sea ice habitat, those findings contributed to the 2008 decision to list the species as threatened under the U.S. Endangered Species Act. Here, we used mark-recapture models to investigate the population dynamics of polar bears in the southern Beaufort Sea from 2001 to 2010, years during which the spatial and temporal extent of summer sea ice generally declined. Low survival from 2004 through 2006 led to a 25-50% decline in abundance. We hypothesize that low survival during this period resulted from (1) unfavorable ice conditions that limited access to prey during multiple seasons; and possibly, (2) low prey abundance. For reasons that are not clear, survival of adults and cubs began to improve in 2007 and abundance was comparatively stable from 2008 to 2010, with ~900 bears in 2010 (90% CI 606-1212). However, survival of subadult bears declined throughout the entire period. Reduced spatial and temporal availability of sea ice is expected to increasingly force population dynamics of polar bears as the climate continues to warm. However, in the short term, our findings suggest that factors other than sea ice can influence survival. A refined understanding of the ecological mechanisms underlying polar bear population dynamics is necessary to improve projections of their future status and facilitate development of management strategies.

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.

Genomic evidence of geographically widespread effect of gene flow from polar bears into brown bears.

Polar bears are an arctic, marine adapted species that is closely related to brown bears. Genome analyses have shown that polar bears are distinct and genetically homogeneous in comparison to brown bears. However, these analyses have also revealed a remarkable episode of polar bear gene flow into the population of brown bears that colonized the Admiralty, Baranof and Chichagof islands (ABC islands) of Alaska. Here, we present an analysis of data from a large panel of polar bear and brown bear genomes that includes brown bears from the ABC islands, the Alaskan mainland and Europe. Our results provide clear evidence that gene flow between the two species had a geographically wide impact, with polar bear DNA found within the genomes of brown bears living both on the ABC islands and in the Alaskan mainland. Intriguingly, while brown bear genomes contain up to 8.8% polar bear ancestry, polar bear genomes appear to be devoid of brown bear ancestry, suggesting the presence of a barrier to gene flow in that direction.

Practice-centred evaluation and the privileging of care in health information technology evaluation.

UNLABELLED: Our contribution, drawn from our experience of the case study provided, is a protocol for practice-centred, participative evaluation of technology in the clinical setting that privileges care. In this context 'practice-centred' evaluation acts as a scalable, coordinating framework for evaluation that recognises health information technology supported care as an achievement that is contingent and ongoing. We argue that if complex programmes of technology-enabled service innovation are understood in terms of their contribution to patient care and supported by participative, capability-building evaluation methodologies, conditions are created for practitioners and patients to realise the potential of technologies and make substantive contributions to the evidence base underpinning health innovation programmes. BACKGROUND: Electronic Patient Records (EPRs) and telemedicine are positioned by policymakers as health information technologies that are integral to achieving improved clinical outcomes and efficiency savings. However, evaluating the extent to which these aims are met poses distinct evaluation challenges, particularly where clinical and cost outcomes form the sole focus of evaluation design. We propose that a practice-centred approach to evaluation - in which those whose day-to-day care practice is altered (or not) by the introduction of new technologies are placed at the centre of evaluation efforts - can complement and in some instances offer advantages over, outcome-centric evaluation models. METHODS: We carried out a regional programme of innovation in renal services where a participative approach was taken to the introduction of new technologies, including: a regional EPR system and a system to support video clinics. An 'action learning' approach was taken to procurement, pre-implementation planning, implementation, ongoing development and evaluation. Participants included clinicians, technology specialists, patients and external academic researchers. Whilst undergoing these activities we asked: how can a practice-centred approach be embedded into evaluation of health information technologies? DISCUSSION: Organising EPR and telemedicine evaluation around predetermined outcome measures alone can be impractical given the complex and contingent nature of such projects. It also limits the extent to which unforeseen outcomes and new capabilities are recognised. Such evaluations often fail to improve understanding of 'when' and 'under what conditions' technology-enabled service improvements are realised, and crucially, how such innovation improves care. SUMMARY: Our contribution, drawn from our experience of the case study provided, is a protocol for practice-centred, participative evaluation of technology in the clinical setting that privileges care. In this context 'practice-centred' evaluation acts as a scalable, coordinating framework for evaluation that recognises health information technology supported care as an achievement that is contingent and ongoing. We argue that if complex programmes of technology-enabled service innovation are understood in terms of their contribution to patient care and supported by participative, capability-building evaluation methodologies, conditions are created for practitioners and patients to realise the potential of technologies and make substantive contributions to the evidence base underpinning health innovation programmes.

Research priorities in spiritual care: an international survey of palliative care researchers and clinicians.

CONTEXT: Spiritual distress, including meaninglessness and hopelessness, is common in advanced disease. Spiritual care is a core component of palliative care, yet often neglected by health care professionals owing to the dearth of robust evidence to guide practice. OBJECTIVES: To determine research priorities of clinicians/researchers and thus inform future research in spiritual care in palliative care. METHODS: An online, cross-sectional, mixed-methods survey was conducted. Respondents were asked whether there is a need for more research in spiritual care, and if so, to select the five most important research priorities from a list of 15 topics. Free-text questions were asked about additional research priorities and respondents' single most important research question, with data analyzed thematically. RESULTS: In total, 971 responses, including 293 from palliative care physicians, 112 from nurses, and 111 from chaplains, were received from 87 countries. Mean age was 48.5 years (standard deviation, 10.7), 64% were women, and 65% were Christian. Fifty-three percent reported their work as "mainly clinical," and less than 2.5% stated that no further research was needed. Integrating quantitative and qualitative data demonstrated three priority areas for research: 1) development and evaluation of conversation models and overcoming barriers to spiritual care in staff attitudes, 2) screening and assessment, and 3) development and evaluation of spiritual care interventions and determining the effectiveness of spiritual care. CONCLUSION: In this first international survey exploring researchers' and clinicians' research priorities in spiritual care, we found international support for research in this domain. Findings provide an evidence base to direct future research and highlight the particular need for methodologically rigorous evaluation studies.

Ecological consequences of sea-ice decline.

After a decade with nine of the lowest arctic sea-ice minima on record, including the historically low minimum in 2012, we synthesize recent developments in the study of ecological responses to sea-ice decline. Sea-ice loss emerges as an important driver of marine and terrestrial ecological dynamics, influencing productivity, species interactions, population mixing, gene flow, and pathogen and disease transmission. Major challenges in the near future include assigning clearer attribution to sea ice as a primary driver of such dynamics, especially in terrestrial systems, and addressing pressures arising from human use of arctic coastal and near-shore areas as sea ice diminishes.

Genomic evidence for island population conversion resolves conflicting theories of polar bear evolution.

Despite extensive genetic analysis, the evolutionary relationship between polar bears (Ursus maritimus) and brown bears (U. arctos) remains unclear. The two most recent comprehensive reports indicate a recent divergence with little subsequent admixture or a much more ancient divergence followed by extensive admixture. At the center of this controversy are the Alaskan ABC Islands brown bears that show evidence of shared ancestry with polar bears. We present an analysis of genome-wide sequence data for seven polar bears, one ABC Islands brown bear, one mainland Alaskan brown bear, and a black bear (U. americanus), plus recently published datasets from other bears. Surprisingly, we find clear evidence for gene flow from polar bears into ABC Islands brown bears but no evidence of gene flow from brown bears into polar bears. Importantly, while polar bears contributed <1% of the autosomal genome of the ABC Islands brown bear, they contributed 6.5% of the X chromosome. The magnitude of sex-biased polar bear ancestry and the clear direction of gene flow suggest a model wherein the enigmatic ABC Island brown bears are the descendants of a polar bear population that was gradually converted into brown bears via male-dominated brown bear admixture. We present a model that reconciles heretofore conflicting genetic observations. We posit that the enigmatic ABC Islands brown bears derive from a population of polar bears likely stranded by the receding ice at the end of the last glacial period. Since then, male brown bear migration onto the island has gradually converted these bears into an admixed population whose phenotype and genotype are principally brown bear, except at mtDNA and X-linked loci. This process of genome erosion and conversion may be a common outcome when climate change or other forces cause a population to become isolated and then overrun by species with which it can hybridize.

What are the toxicological effects of mercury in Arctic biota?

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 of mercury recorded in brain tissue with associated signs of neurochemical effects. Evidence of increasing concentrations in mercury in some biota in Arctic Canada and Greenland is therefore a concern with respect to ecosystem health.

Age and sex composition of seals killed by polar bears in the eastern Beaufort Sea.

BACKGROUND: Polar bears (Ursus maritimus) of the Beaufort Sea enter hyperphagia in spring and gain fat reserves to survive periods of low prey availability. We collected information on seals killed by polar bears (n=650) and hunting attempts on ringed seal (Pusa hispida) lairs (n=1396) observed from a helicopter during polar bear mark-recapture studies in the eastern Beaufort Sea in spring in 1985-2011. We investigated how temporal shifts in ringed seal reproduction affect kill composition and the intraspecific vulnerabilities of ringed seals to polar bear predation. PRINCIPAL FINDINGS: Polar bears primarily preyed on ringed seals (90.2%) while bearded seals (Erignathus barbatus) only comprised 9.8% of the kills, but 33% of the biomass. Adults comprised 43.6% (150/344) of the ringed seals killed, while their pups comprised 38.4% (132/344). Juvenile ringed seals were killed at the lowest proportion, comprising 18.0% (62/344) of the ringed seal kills. The proportion of ringed seal pups was highest between 2007-2011, in association with high ringed seal productivity. Half of the adult ringed seal kills were ≥ 21 years (60/121), and kill rates of adults increased following the peak of parturition. Determination of sex from DNA revealed that polar bears killed adult male and adult female ringed seals equally (0.50, n=78). The number of hunting attempts at ringed seal subnivean lair sites was positively correlated with the number of pup kills (r(2) =0.30, P=0.04), but was not correlated with the number of adult kills (P=0.37). CONCLUSIONS/SIGNIFICANCE: Results are consistent with decadal trends in ringed seal productivity, with low numbers of pups killed by polar bears in spring in years of low pup productivity, and conversely when pup productivity was high. Vulnerability of adult ringed seals to predation increased in relation to reproductive activities and age, but not gender.

Effects of climate warming on polar bears: a review of the evidence.

Climate warming is causing unidirectional changes to annual patterns of sea ice distribution, structure, and freeze-up. We summarize evidence that documents how loss of sea ice, the primary habitat of polar bears (Ursus maritimus), negatively affects their long-term survival. To maintain viable subpopulations, polar bears depend on sea ice as a platform from which to hunt seals for long enough each year to accumulate sufficient energy (fat) to survive periods when seals are unavailable. Less time to access to prey, because of progressively earlier breakup in spring, when newly weaned ringed seal (Pusa hispida) young are available, results in longer periods of fasting, lower body condition, decreased access to denning areas, fewer and smaller cubs, lower survival of cubs as well as bears of other age classes and, finally, subpopulation decline toward eventual extirpation. The chronology of climate-driven changes will vary between subpopulations, with quantifiable negative effects being documented first in the more southerly subpopulations, such as those in Hudson Bay or the southern Beaufort Sea. As the bears' body condition declines, more seek alternate food resources so the frequency of conflicts between bears and humans increases. In the most northerly areas, thick multiyear ice, through which little light penetrates to stimulate biological growth on the underside, will be replaced by annual ice, which facilitates greater productivity and may create habitat more favorable to polar bears over continental shelf areas in the short term. If the climate continues to warm and eliminate sea ice as predicted, polar bears will largely disappear from the southern portions of their range by mid-century. They may persist in the northern Canadian Arctic Islands and northern Greenland for the foreseeable future, but their long-term viability, with a much reduced global population size in a remnant of their former range, is uncertain.