Eric V. Regehr

Reduced body size and cub recruitment in polar bears associated with sea ice decline

Rates of reproduction and survival are dependent upon adequate body size and condition of individuals. Declines in size and condition have provided early indicators of population decline in polar bears (Ursus maritimus) near the southern extreme of their range. We tested whether patterns in body size, condition, and cub recruitment of polar bears in the southern Beaufort Sea of Alaska were related to the availability of preferred sea ice habitats and whether these measures and habitat availability exhibited trends over time, between 1982 and 2006. The mean skull size and body length of all polar bears over three years of age declined over time, corresponding with long-term declines in the spatial and temporal availability of sea ice habitat. Body size of young, growing bears declined over time and was smaller after years when sea ice availability was reduced. Reduced litter mass and numbers of yearlings per female following years with lower availability of optimal sea ice habitat, suggest reduced reproductive output and juvenile survival. These results, based on analysis of a long-term data set, suggest that declining sea ice is associated with nutritional limitations that reduced body size and reproduction in this population.

Arctic marine mammal population status, sea ice habitat loss, and conservation recommendations for the 21st century

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

Variation in the response of an Arctic top predator experiencing habitat loss: feeding and reproductive ecology of two polar bear populations

Polar bears (Ursus maritimus) have experienced substantial changes in the seasonal availability of sea ice habitat in parts of their range, including the Beaufort, Chukchi, and Bering Seas. In this study, we compared the body size, condition, and recruitment of polar bears captured in the Chukchi and Bering Seas (CS) between two periods (1986-1994 and 2008-2011) when declines in sea ice habitat occurred. In addition, we compared metrics for the CS population 2008-2011 with those of the adjacent southern Beaufort Sea (SB) population where loss in sea ice habitat has been associated with declines in body condition, size, recruitment, and survival. We evaluated how variation in body condition and recruitment were related to feeding ecology. Comparing habitat conditions between populations, there were twice as many reduced ice days over continental shelf waters per year during 2008-2011 in the SB than in the CS. CS polar bears were larger and in better condition, and appeared to have higher reproduction than SB bears. Although SB and CS bears had similar diets, twice as many bears were fasting in spring in the SB than in the CS. Between 1986-1994 and 2008-2011, body size, condition, and recruitment indices in the CS were not reduced despite a 44-day increase in the number of reduced ice days. Bears in the CS exhibited large body size, good body condition, and high indices of recruitment compared to most other populations measured to date. Higher biological productivity and prey availability in the CS relative to the SB, and a shorter recent history of reduced sea ice habitat, may explain the maintenance of condition and recruitment of CS bears. Geographic differences in the response of polar bears to climate change are relevant to range-wide forecasts for this and other ice-dependent species.

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.

Polar bear population status in the northern Beaufort Sea, Canada, 1971—2006

Polar bears (Ursus maritimus) of the northern Beaufort Sea (NB) population occur on the perimeter of the polar basin adjacent to the northwestern islands of the Canadian Arctic Archipelago. Sea ice converges on the islands through most of the year. We used open-population capture-recapture models to estimate population size and vital rates of polar bears between 1971 and 2006 to: (1) assess relationships between survival, sex and age, and time period; (2) evaluate the long-term importance of sea ice quality and availability in relation to climate warming; and (3) note future management and conservation concerns. The highest-ranking models suggested that survival of polar bears varied by age class and with changes in the sea ice habitat. Model-averaged estimates of survival (which include harvest mortality) for senescent adults ranged from 0.37 to 0.62, from 0.22 to 0.68 for cubs of the year (COY) and yearlings, and from 0.77 to 0.92 for 2-4 year-olds and adults. Horvtiz-Thompson (HT) estimates of population size were not significantly different among the decades of our study. The population size estimated for the 2000s was 980 +/- 155 (mean and 95% CI). These estimates apply primarily to that segment of the NB population residing west and south of Banks Island. The NB polar bear population appears to have been stable or possibly increasing slightly during the period of our study. This suggests that ice conditions have remained suitable and similar for feeding in summer and fall during most years and that the traditional and legal Inuvialuit harvest has not exceeded sustainable levels. However, the amount of ice remaining in the study area at the end of summer, and the proportion that continues to lie over the biologically productive continental shelf (< 300 m water depth) has declined over the 35-year period of this study. If the climate continues to warm as predicted, we predict that the polar bear population in the northern Beaufort Sea will eventually decline. Management and conservation practices for polar bears in relation to both aboriginal harvesting and offshore industrial activity will need to adapt.

Summer declines in activity and body temperature offer polar bears limited energy savings

Not that unusual after all As polar ice recedes, polar bears are facing a changed habitat with reduced summer foraging opportunities. It has been hypothesized that they might be able to resist summer food shortages by reducing their metabolic needs in a sort of “walking hibernation.” Whiteman et al. monitored energy expenditure in polar bears both on and off the ice and found energy reductions, but that these were more akin to normal mammalian fasting levels. Thus, it appears that polar bears have no energetic protections against reduced summer food supplies and will face increasing starvation threats if summer foraging habitats continue to decline. Science, this issue p. 295 Polar bears show no inherent energetic resistance to summer food shortages. Polar bears (Ursus maritimus) summer on the sea ice or, where it melts, on shore. Although the physiology of “ice” bears in summer is unknown, “shore” bears purportedly minimize energy losses by entering a hibernation-like state when deprived of food. Such a strategy could partially compensate for the loss of on-ice foraging opportunities caused by climate change. However, here we report gradual, moderate declines in activity and body temperature of both shore and ice bears in summer, resembling energy expenditures typical of fasting, nonhibernating mammals. Also, we found that to avoid unsustainable heat loss while swimming, bears employed unusual heterothermy of the body core. Thus, although well adapted to seasonal ice melt, polar bears appear susceptible to deleterious declines in body condition during the lengthening period of summer food deprivation.

Increased Land Use by Chukchi Sea Polar Bears in Relation to Changing Sea Ice Conditions

Recent observations suggest that polar bears (Ursus maritimus) are increasingly using land habitats in some parts of their range, where they have minimal access to their preferred prey, likely in response to loss of their sea ice habitat associated with climatic warming. We used location data from female polar bears fit with satellite radio collars to compare land use patterns in the Chukchi Sea between two periods (1986–1995 and 2008–2013) when substantial summer sea-ice loss occurred. In both time periods, polar bears predominantly occupied sea-ice, although land was used during the summer sea-ice retreat and during the winter for maternal denning. However, the proportion of bears on land for > 7 days between August and October increased between the two periods from 20.0% to 38.9%, and the average duration on land increased by 30 days. The majority of bears that used land in the summer and for denning came to Wrangel and Herald Islands (Russia), highlighting the importance of these northernmost land habitats to Chukchi Sea polar bears. Where bears summered and denned, and how long they spent there, was related to the timing and duration of sea ice retreat. Our results are consistent with other studies supporting increased land use as a common response of polar bears to sea-ice loss. Implications of increased land use for Chukchi Sea polar bears are unclear, because a recent study observed no change in body condition or reproductive indices between the two periods considered here. This result suggests that the ecology of this region may provide a degree of resilience to sea ice loss. However, projections of continued sea ice loss suggest that polar bears in the Chukchi Sea and other parts of the Arctic may increasingly use land habitats in the future, which has the potential to increase nutritional stress and human-polar bear interactions.

Polar bear population status in southern Hudson Bay, Canada

5 Introduction 5 Methods and Materials 3 Study Area 3 Capture Methods 4 Capture-recapture analysis 5 Goodness-of-fit 6 Covariates 6 Model selection 8 Parameter estimates 8 Results 9 Captures 9 Goodness of fit 9 Model selection 9 Survival Estimates 10 Abundance Estimates 11 Discussion 11 Interpretation of Parameter Estimates 11 Survival 12 Population size and trend 12 Future trend 14 Acknowledgments 14 References Cited 15

Identifying polar bear resource selection patterns to inform offshore development in a dynamic and changing Arctic

Although sea ice loss is the primary threat to polar bears (Ursus maritimus), little can be done to mitigate its effects without global efforts to reduce greenhouse gas emissions. Other factors, however, could exacerbate the impacts of sea ice loss on polar bears, such as exposure to increased industrial activity. The Arctic Ocean has enormous oil and gas potential, and its development is expected to increase in the coming decades. Estimates of polar bear resource selection will inform managers how bears use areas slated for oil development and to help guide conservation planning. We estimated temporally-varying resource selection patterns for non-denning adult female polar bears in the Chukchi Sea population (2008–2012) at two scales (i.e., home range and weekly steps) to identify factors predictive of polar bear use throughout the year, before any offshore development. From the best models at each scale, we estimated scale-integrated resource selection functions to predict polar bear space use across th...