Gregory W. Thiemann

POLAR BEAR DIETS AND ARCTIC MARINE FOOD WEBS: INSIGHTS FROM FATTY ACID ANALYSIS

We used quantitative fatty acid signature analysis (QFASA) to examine the diets of 1738 individual polar bears (Ursus maritimus) sampled across the Canadian Arctic over a 30-year span. Polar bear foraging varied over large and small spatial and temporal scales, and between demographic groups. Diets in every subpopulation were dominated by ringed seals (Phoca hispida) and, in the eastern Arctic, secondarily by harp seals (Pagophilus groenlandica). Beluga whales (Delphinapterus leucas) were an important food source for bears in the High Arctic, which is consistent with previous anecdotal reports. Foraging patterns were most similar among neighboring subpopulations with similar prey assemblages, but also differed geographically within Western Hudson Bay. The sexual size dimorphism of polar bears had an important effect on foraging, as large bearded seals (Erignathus barbatus) and walruses (Odobenus rosmarus) were consumed most often by older, male bears, whereas ringed seals and, where available, harbor seals (P. vitulina) were most important to younger age classes. Larger, older bears also had the greatest dietary diversity, apparently because of their ability to include larger-bodied prey. During spring and summer, polar bears in some areas increased predation on migratory harp seals and beluga whales. In Western Hudson Bay, bearded seal consumption declined between 1995 and 2001 for both male and female bears and continued to decline among females up to the most recent sampling (2004). Ringed seal consumption in Western Hudson Bay increased between 1998 and 2001, perhaps in response to increased ringed seal productivity, but was not significantly affected by date of sea-ice breakup. Overall, our data indicate that polar bears are capable of opportunistically altering their foraging to take advantage of locally abundant prey, or to some degree compensating for a decline in a dominant prey species. However, in other areas polar bears are dependent on the availability of ringed and bearded seals. Recent population data suggest that polar bears with the most specialized diets may be most vulnerable to climate-related changes in ice conditions. The results of this large-scale, ecosystem-based study indicate a complex relationship between sea-ice conditions, prey population dynamics, and polar bear foraging.

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.

Migration phenology and seasonal fidelity of an Arctic marine predator in relation to sea ice dynamics

Understanding how seasonal environmental conditions affect the timing and distribution of synchronized animal movement patterns is a central issue in animal ecology. Migration, a behavioural adaptation to seasonal environmental fluctuations, is a fundamental part of the life history of numerous species. However, global climate change can alter the spatiotemporal distribution of resources and thus affect the seasonal movement patterns of migratory animals. We examined sea ice dynamics relative to migration patterns and seasonal geographical fidelity of an Arctic marine predator, the polar bear (Ursus maritimus). Polar bear movement patterns were quantified using satellite-linked telemetry data collected from collars deployed between 1991-1997 and 2004-2009. We showed that specific sea ice characteristics can predict the timing of seasonal polar bear migration on and off terrestrial refugia. In addition, fidelity to specific onshore regions during the ice-free period was predicted by the spatial pattern of sea ice break-up but not by the timing of break-up. The timing of migration showed a trend towards earlier arrival of polar bears on shore and later departure from land, which has been driven by climate-induced declines in the availability of sea ice. Changes to the timing of migration have resulted in polar bears spending progressively longer periods of time on land without access to sea ice and their marine mammal prey. The links between increased atmospheric temperatures, sea ice dynamics, and the migratory behaviour of an ice-dependent species emphasizes the importance of quantifying and monitoring relationships between migratory wildlife and environmental cues that may be altered by climate change.

Quantitative Support for a Subjective Fatness Index for Immobilized Polar Bears

Abstract Adequate stores of body fat are essential for survival and reproduction of polar bears (Ursus maritimus). However, polar bear body fat levels can be difficult to quantify in the field. For >30 years, biologists have subjectively estimated relative fatness of immobilized polar bears by assigning individuals a rating from 1 to 5, with 1 being leanest and 5 most obese. Although previous studies suggested this fatness index (FI) rating accurately reflects large-scale differences in body condition, its relationship to more quantitative measures of condition has not been explored. We compared the FI rating of individual polar bears in western Hudson Bay and the Beaufort Sea to 2 quantitative measures of body condition: the Quetelet Index (ratio of mass to length2) and the relative lipid content of adipose tissue. We found a significant relationship between FI rating and both Quetelet Index values and adipose lipid content. Our data demonstrate that the FI rating accurately reflects overall body condition, regardless of polar bear age, sex, or nutritional phase. We suggest that continued field use of the FI rating could provide valuable information on ecological effects of large-scale environmental change on polar bear populations.

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.

Distance measures and optimization spaces in quantitative fatty acid signature analysis

Quantitative fatty acid signature analysis has become an important method of diet estimation in ecology, especially marine ecology. Controlled feeding trials to validate the method and estimate the calibration coefficients necessary to account for differential metabolism of individual fatty acids have been conducted with several species from diverse taxa. However, research into potential refinements of the estimation method has been limited. We compared the performance of the original method of estimating diet composition with that of five variants based on different combinations of distance measures and calibration-coefficient transformations between prey and predator fatty acid signature spaces. Fatty acid signatures of pseudopredators were constructed using known diet mixtures of two prey data sets previously used to estimate the diets of polar bears Ursus maritimus and gray seals Halichoerus grypus, and their diets were then estimated using all six variants. In addition, previously published diets of Chukchi Sea polar bears were re-estimated using all six methods. Our findings reveal that the selection of an estimation method can meaningfully influence estimates of diet composition. Among the pseudopredator results, which allowed evaluation of bias and precision, differences in estimator performance were rarely large, and no one estimator was universally preferred, although estimators based on the Aitchison distance measure tended to have modestly superior properties compared to estimators based on the Kullback–Leibler distance measure. However, greater differences were observed among estimated polar bear diets, most likely due to differential estimator sensitivity to assumption violations. Our results, particularly the polar bear example, suggest that additional research into estimator performance and model diagnostics is warranted.

Geographical Area and Life History Traits Influence Diet in an Arctic Marine Predator.

Global changes are thought to affect most Arctic species, yet some populations are more at risk. Today, the Barents Sea ecoregion is suffering the strongest sea ice retreat ever measured; and these changes are suspected to modify food access and thus diet of several species. Biochemical diet tracers enable investigation of diet in species such as polar bears (Ursus maritimus). We examined individual diet variation of female polar bears in Svalbard, Norway, and related it to year, season (spring and autumn), sampling area and breeding status (solitary, with cubs of the year or yearlings). Sampling areas were split according to their ice cover: North-West (less sea ice cover), South-East (larger amplitude in sea ice extent) and North-East/South-West (NESW) as bears from that zone are more mobile among all regions of Svalbard. We measured fatty acid (FA) composition in adipose tissue and carbon (δ13C) and nitrogen (δ15N) stable isotopes in plasma and red blood cells. Females feeding in the North-West area had lower δ15N values than those from the NESW. In South-East females, δ13C values were lower in autumn compared to spring and females seemed less selective in their diet as depicted by large variances in stable isotope values. Considering the differences in FA composition and stable isotope values, we suggest that females from the North-West and South-East could ingest a higher proportion of avian prey. With regard to breeding status, solitary females had higher δ15N values and smaller variance in their stable isotopic values than females with cubs, suggesting that solitary females were more selective and prey on higher trophic level species (i.e. seals). Overall, our results indicate that prey availability for Svalbard polar bears varies according to geographical area and prey selectivity differs according to breeding status. Our findings suggest that complex changes in sea ice and prey availability will interact to affect Svalbard polar bear feeding patterns and associated nutrition.

Assessing the robustness of quantitative fatty acid signature analysis to assumption violations

Summary Knowledge of animal diets can provide important insights into life history and ecology, relationships among species in a community and potential response to ecosystem change or perturbation. Quantitative fatty acid signature analysis (QFASA) is a method of estimating diets from data on the composition, or signature, of fatty acids stored in adipose tissue. Given data on signatures of potential prey, a predator diet is estimated by minimizing the distance between its signature and a mixture of prey signatures. Calibration coefficients, constants derived from feeding trials, are used to account for differential metabolism of individual fatty acids. QFASA has been widely applied since its introduction and several variants of the original estimator have appeared in the literature. However, work to compare the statistical properties of QFASA estimators has been limited. One important characteristic of an estimator is its robustness to violations of model assumptions. The primary assumptions of QFASA are that prey signature data contain representatives of all prey types consumed and the calibration coefficients are known without error. We investigated the robustness of two QFASA estimators to a range of violations of these assumptions using computer simulation and recorded the resulting bias in diet estimates. We found that the Aitchison distance measure was most robust to errors in the calibration coefficients. Conversely, the Kullback–Leibler distance measure was most robust to the consumption of prey without representation in the prey signature data. In most QFASA applications, investigators will generally have some knowledge of the prey available to predators and be able to assess the completeness of prey signature data and sample additional prey as necessary. Conversely, because calibration coefficients are derived from feeding trials with captive animals and their values may be sensitive to consumer physiology and nutritional status, their applicability to free-ranging animals is difficult to establish. We therefore recommend that investigators first make any improvements to the prey signature data that seem warranted and then base estimation on the Aitchison distance measure, as it appears to minimize risk from violations of the assumption that is most difficult to verify.

Effects of chemical immobilization on the movement rates of free-ranging polar bears

Abstract The capture and handling of free-ranging animals is an important tool for wildlife research, conservation, and management. However, live capture may expose individual animals to risk of injury, impairment, or mortality. The polar bear (Ursus maritimus) is a species of conservation concern throughout its range and physical mark–recapture techniques have formed the basis of polar bear research and harvest management for decades. We examined movement patterns of polar bears postcapture to measure their recovery from chemical immobilization and determine whether captured bears experienced prolonged effects that would affect individual fitness. Adult female (n = 61) and juvenile (n = 13) polar bears in 3 Canadian subpopulations were captured during the course of other studies using a combination of tiletamine hydrochloride and zolazepam hydrochloride delivered via remote injection from a helicopter. Bears were fitted with satellite-linked global positioning system collars and we used 3 individual-based metrics to assess their recovery from immobilization: time to move 50 m; time to move 100 m; and time to reach a baseline movement rate threshold (km/day) derived from each individuals movements in a fully recovered state (i.e., 30–60 days postcapture). There were no differences in recovery rate metrics across years, age classes, or between females with cubs of different ages. When compared across subpopulations, only the time to move 50 m differed, being shortest in the southern Beaufort Sea. Bears captured on land during the ice-free period in western Hudson Bay and Foxe Basin were more variable in their response to capture than were those handled on the sea ice of the Beaufort Sea, but in all 3 areas, bears showed gradual increases in movement rates. Movement rates indicative of recovery were often reached 48 h after capture and 51 (69%) of 74 bears appeared to be fully recovered in ≤3 days. Consistent with preliminary work on chemical immobilization of polar bears, there was no relationship between drug dose and rate of recovery. Our results indicated that polar bears captured in different locations, seasons, and life-history stages recovered predictably from chemical immobilization in a time frame that is unlikely to affect individual fitness.

Sea ice-associated decline in body condition leads to increased concentrations of lipophilic pollutants in polar bears (Ursus maritimus) from Svalbard, Norway

Global climate changes are magnified in the Arctic and are having an especially dramatic effect on the spatial and temporal distribution and the thickness traits of sea ice. Decline of Arctic sea ice may lead to qualitative and/or quantitative changes in diet and reduced body condition (i.e. adipose tissue stores) of ice-associated apex predators such as polar bears (Ursus maritimus). This may further affect their tissue concentrations of lipophilic pollutants. We determined how variations in adipose tissue stores associated to both breeding status and spatial changes in sea ice conditions and diet influence concentrations and biotransformation of lipophilic persistent organic pollutants (POPs). We collected 112 blood and fat samples from female polar bears (Ursus maritimus) of different breeding status (alone, with cubs of the year, or with yearlings) during two seasons (April and September) in 2012 and 2013 at three locations of Svalbard, Norway, with contrasted sea ice conditions. We inferred diet from nitrogen and carbon stable isotope ratios in red blood cells and fatty acid composition in adipose tissue. Relative to diet, body condition, which was negatively related to sea ice extent at both temporal and spatial scales, was the most important predictor for concentrations of POPs in plasma and fat, whereas diet showed a minor influence. Additionally, fatter females were more efficient at biotransforming PCBs than were leaner ones. Breeding status influenced the concentrations of less lipophilic compounds such as β-hexachlorocyclohexane, which were lower in females with yearlings, probably due to excretion into milk and subsequent offloading to young. In conclusion, our results indicate that declining sea ice indirectly leads to increased concentrations of lipophilic pollutants in polar bears mediated through reduced feeding opportunities and declining body condition rather than changes in diet composition.