Simultaneous estimation of the temporal and spatial extent of animal migration using step lengths and turning angles.

BACKGROUND: Animals of many different species, trophic levels, and life history strategies migrate, and the improvement of animal tracking technology allows ecologists to collect increasing amounts of detailed data on these movements. Understanding when animals migrate is important for managing their populations, but is still difficult despite modelling advancements. METHODS: We designed a model that parametrically estimates the timing of migration from animal tracking data. Our model identifies the beginning and end of migratory movements as signaled by change-points in step length and turning angle distributions. To this end, we can also use the model to estimate how an animal's movement changes when it begins migrating. In addition to a thorough simulation analysis, we tested our model on three datasets: migratory ferruginous hawks (Buteo regalis) in the Great Plains, barren-ground caribou (Rangifer tarandus groenlandicus) in northern Canada, and non-migratory brown bears (Ursus arctos) from the Canadian Arctic. RESULTS: Our simulation analysis suggests that our model is most useful for datasets where an increase in movement speed or directional autocorrelation is clearly detectable. We estimated the beginning and end of migration in caribou and hawks to the nearest day, while confirming a lack of migratory behaviour in the brown bears. In addition to estimating when caribou and ferruginous hawks migrated, our model also identified differences in how they migrated; ferruginous hawks achieved efficient migrations by drastically increasing their movement rates while caribou migration was achieved through significant increases in directional persistence. CONCLUSIONS: Our approach is applicable to many animal movement studies and includes parameters that can facilitate comparison between different species or datasets. We hope that rigorous assessment of migration metrics will aid understanding of both how and why animals move.

Survival improvements of marine mammals in zoological institutions mirror historical advances in human longevity.

An intense public debate has fuelled governmental bans on marine mammals held in zoological institutions. The debate rests on the assumption that survival in zoological institutions has been and remains lower than in the wild, albeit the scientific evidence in support of this notion is equivocal. Here, we used statistical methods previously applied to assess historical improvements in human lifespan and data on 8864 individuals of four marine mammal species (harbour seal, Phoca vitulina; California sea lion, Zalophus californianus; polar bear, Ursus maritimus; common bottlenose dolphin, Tursiops truncatus) held in zoos from 1829 to 2020. We found that life expectancy increased up to 3.40 times, and first-year mortality declined up to 31%, during the last century in zoos. Moreover, the life expectancy of animals in zoos is currently 1.65-3.55 times longer than their wild counterparts. Like humans, these improvements have occurred concurrently with advances in management practices, crucial for population welfare. Science-based decisions will help effective legislative changes and ensure better implementation of animal care.

Diet energy density estimated from isotopes in predator hair associated with survival, habitat, and population dynamics.

Sea ice loss is fundamentally altering the Arctic marine environment. Yet there is a paucity of data on the adaptability of food webs to ecosystem change, including predator-prey interactions. Polar bears (Ursus maritimus) are an important subsistence resource for Indigenous people and an apex predator that relies entirely on the under-ice food web to meet its energy needs. In this study, we assessed whether polar bears maintained dietary energy density by prey switching in response to spatiotemporal variation in prey availability. We compared the macronutrient composition of diets inferred from stable carbon and nitrogen isotopes in polar bear guard hair (primarily representing summer/fall diet) during periods when bears had low and high survival (2004-2016), between bears that summered on land versus pack ice, and between bears occupying different regions of the Alaskan and Canadian Beaufort Sea. Polar bears consumed diets with lower energy density during periods of low survival, suggesting that concurrent increased dietary proportions of beluga whales (Delphinapterus leucas) did not offset reduced proportions of ringed seals (Pusa hispida). Diets with the lowest energy density and proportions from ringed seal blubber were consumed by bears in the western Beaufort Sea (Alaska) during a period when polar bear abundance declined. Intake required to meet energy requirements of an average free-ranging adult female polar bear was 2.1 kg/day on diets consumed during years with high survival but rose to 3.0 kg/day when survival was low. Although bears that summered onshore in the Alaskan Beaufort Sea had higher-fat diets than bears that summered on the pack ice, access to the remains of subsistence-harvested bowhead whales (Balaena mysticetus) contributed little to improving diet energy density. Because most bears in this region remain with the sea ice year round, prey switching and consumption of whale carcasses onshore appear insufficient to augment diets when availability of their primary prey, ringed seals, is reduced. Our results show that a strong predator-prey relationship between polar bears and ringed seals continues in the Beaufort Sea. The method of estimating dietary blubber using predator hair, demonstrated here, provides a new metric to monitor predator-prey relationships that affect individual health and population demographics.

Drivers of polar bear behavior and the possible effects of prey availability on foraging strategy.

BACKGROUND: Change in behavior is one of the earliest responses to variation in habitat suitability. It is therefore important to understand the conditions that promote different behaviors, particularly in areas undergoing environmental change. Animal movement is tightly linked to behavior and remote tracking can be used to study ethology when direct observation is not possible. METHODS: We used movement data from 14 polar bears (Ursus maritimus) in Hudson Bay, Canada, during the foraging season (January-June), when bears inhabit the sea ice. We developed an error-tolerant method to correct for sea ice drift in tracking data. Next, we used hidden Markov models with movement and orientation relative to wind to study three behaviors (stationary, area-restricted search, and olfactory search) and examine effects of 11 covariates on behavior. RESULTS: Polar bears spent approximately 47% of their time in the stationary drift state, 29% in olfactory search, and 24% in area-restricted search. High energy behaviors occurred later in the day (around 20:00) compared to other populations. Second, olfactory search increased as the season progressed, which may reflect a shift in foraging strategy from still-hunting to active search linked to a shift in seal availability (i.e., increase in haul-outs from winter to the spring pupping and molting seasons). Last, we found spatial patterns of distribution linked to season, ice concentration, and bear age that may be tied to habitat quality and competitive exclusion. CONCLUSIONS: Our observations were generally consistent with predictions of the marginal value theorem, and differences between our findings and other populations could be explained by regional or temporal variation in resource availability. Our novel movement analyses and finding can help identify periods, regions, and conditions of critical habitat.

Time-dependent memory and individual variation in Arctic brown bears (Ursus arctos).

BACKGROUND: Animal movement modelling provides unique insight about how animals perceive their landscape and how this perception may influence space use. When coupled with data describing an animal's environment, ecologists can fit statistical models to location data to describe how spatial memory informs movement. METHODS: We performed such an analysis on a population of brown bears (Ursus arctos) in the Canadian Arctic using a model incorporating time-dependent spatial memory patterns. Brown bear populations in the Arctic lie on the periphery of the species' range, and as a result endure harsh environmental conditions. In this kind of environment, effective use of memory to inform movement strategies could spell the difference between survival and mortality. RESULTS: The model we fit tests four alternate hypotheses (some incorporating memory; some not) against each other, and we found a high degree of individual variation in how brown bears used memory. We found that 71% (15 of 21) of the bears used complex, time-dependent spatial memory to inform their movement decisions. CONCLUSIONS: These results, coupled with existing knowledge on individual variation in the population, highlight the diversity of foraging strategies for Arctic brown bears while also displaying the inference that can be drawn from this innovative movement model.

Energy-based step selection analysis: Modelling the energetic drivers of animal movement and habitat use.

The energetic gains from foraging and costs of movement are expected to be key drivers of animal decision-making, as their balance is a large determinant of body condition and survival. This fundamental perspective is often missing from habitat selection studies, which mainly describe correlations between space use and environmental features, rather than the mechanisms behind these correlations. To address this gap, we present a novel parameterisation of step selection functions (SSFs), that we term the energy selection function (ESF). In this model, the likelihood of an animal selecting a movement step depends directly on the corresponding energetic gains and costs, and we can therefore assess how moving animals choose habitat based on energetic considerations. The ESF retains the mathematical convenience and practicality of other SSFs and can be quickly fitted using standard software. In this article, we outline a workflow, from data gathering to statistical analysis, and use a case study of polar bears Ursus maritimus to demonstrate application of the model. We explain how defining gains and costs at the scale of the movement step allows us to include information about resource distribution, landscape resistance and movement patterns. We further demonstrate this process with a case study of polar bears and show how the parameters can be interpreted in terms of selection for energetic gains and against energetic costs. The ESF is a flexible framework that combines the energetic consequences of both movement and resource selection, thus incorporating a key mechanism into habitat selection analysis. Further, because it is based on familiar habitat selection models, the ESF is widely applicable to any study system where energetic gains and costs can be derived, and has immense potential for methodological extensions.

A risk assessment review of mercury exposure in Arctic marine and terrestrial mammals.

There has been a considerable number of reports on Hg concentrations in Arctic mammals since the last Arctic Monitoring and Assessment Programme (AMAP) effort to review biological effects of the exposure to mercury (Hg) in Arctic biota in 2010 and 2018. Here, we provide an update on the state of the knowledge of health risk associated with Hg concentrations in Arctic marine and terrestrial mammal species. Using available population-specific data post-2000, our ultimate goal is to provide an updated evidence-based estimate of the risk for adverse health effects from Hg exposure in Arctic mammal species at the individual and population level. Tissue residues of Hg in 13 species across the Arctic were classified into five risk categories (from No risk to Severe risk) based on critical tissue concentrations derived from experimental studies on harp seals and mink. Exposure to Hg lead to low or no risk for health effects in most populations of marine and terrestrial mammals, however, subpopulations of polar bears, pilot whales, narwhals, beluga and hooded seals are highly exposed in geographic hotspots raising concern for Hg-induced toxicological effects. About 6% of a total of 3500 individuals, across different marine mammal species, age groups and regions, are at high or severe risk of health effects from Hg exposure. The corresponding figure for the 12 terrestrial species, regions and age groups was as low as 0.3% of a total of 731 individuals analyzed for their Hg loads. Temporal analyses indicated that the proportion of polar bears at low or moderate risk has increased in East/West Greenland and Western Hudson Bay, respectively. However, there remain numerous knowledge gaps to improve risk assessments of Hg exposure in Arctic mammalian species, including the establishment of improved concentration thresholds and upscaling to the assessment of population-level effects.

Sea ice reduction drives genetic differentiation among Barents Sea polar bears.

Loss of Arctic sea ice owing to climate change is predicted to reduce both genetic diversity and gene flow in ice-dependent species, with potentially negative consequences for their long-term viability. Here, we tested for the population-genetic impacts of reduced sea ice cover on the polar bear (Ursus maritimus) sampled across two decades (1995-2016) from the Svalbard Archipelago, Norway, an area that is affected by rapid sea ice loss in the Arctic Barents Sea. We analysed genetic variation at 22 microsatellite loci for 626 polar bears from four sampling areas within the archipelago. Our results revealed a 3-10% loss of genetic diversity across the study period, accompanied by a near 200% increase in genetic differentiation across regions. These effects may best be explained by a decrease in gene flow caused by habitat fragmentation owing to the loss of sea ice coverage, resulting in increased inbreeding of local polar bears within the focal sampling areas in the Svalbard Archipelago. This study illustrates the importance of genetic monitoring for developing adaptive management strategies for polar bears and other ice-dependent species.

Comparing sea ice habitat fragmentation metrics using integrated step selection analysis.

Habitat fragmentation occurs when continuous habitat gets broken up as a result of ecosystem change. While commonly studied in terrestrial ecosystems, Arctic sea ice ecosystems also experience fragmentation, but are rarely studied in this context. Most fragmentation analyses are conducted using patch-based metrics, which are potentially less suitable for sea ice that has gradual changes between sea ice cover, than distinct "long-term" patches. Using an integrated step selection analysis, we compared the descriptive power of a patch-based metric to a more novel metric, the variation in local spatial autocorrelation over time. We used satellite telemetry data from 39 adult female polar bears (Ursus maritimus) in Hudson Bay to examine their sea ice habitat using Advanced Microwave Scanning Radiometer 2 data during sea ice breakup in May through July from 2013-2018. Spatial autocorrelation resulted in better model fits across 64% of individuals, although both metrics were more effective in describing movement patterns than habitat selection. Variation in local spatial autocorrelation allows for the visualization of sea ice habitat at complex spatial and temporal scales, condensing a targeted time period of habitat that would otherwise have to be analyzed daily.

Spatial and temporal variability in ringed seal (Pusa hispida) stable isotopes in the Beaufort Sea.

Arctic ecosystem dynamics are shifting in response to warming temperatures and sea ice loss. Such ecosystems may be monitored by examining the diet of upper trophic level species, which varies with prey availability. To assess interannual variation in the Beaufort Sea ecosystem, we examined spatial and temporal trends in ringed seal (Pusa hispida) δ13C and δ15N in claw growth layers grown from 1964 to 2011. Stable isotopes were correlated with climate indices, environmental conditions, seal population productivity, and geographic location. Sex and age did not influence stable isotopes. Enriched 13C was linked to cyclonic circulation regimes, seal productivity, and westward sampling locations. Higher δ15N was linked to lower sea surface temperatures, a higher percentage of pups in the subsistence harvest, and sample locations that were eastward and further from shore. From the 1960s to 2000s, ringed seal niche width expanded, suggesting a diversification of diet due to expansion of prey and/or seal space use. Overall, trends in ringed seal stable isotopes indicate changes within the Beaufort Sea ecosystem affected by water temperatures and circulation regimes. We suggest that continued monitoring of upper trophic level species will yield insights into changing ecosystem structure with climate change.

Two Decades of Mercury Concentrations in Barents Sea Polar Bears (Ursus maritimus) in Relation to Dietary Carbon, Sulfur, and Nitrogen.

Temporal trends of total mercury (THg) were examined in female polar bear (Ursus maritimus) hair (n = 199) from the Barents Sea in 1995-2016. In addition, hair values of stable isotopes (n = 190-197) of carbon (δ13C), sulfur (δ34S), and nitrogen (δ15N) and information on breeding status, body condition, and age were obtained. Stable isotope values of carbon and sulfur reflect dietary source (e.g., marine vs terrestrial) and the nitrogen trophic level. Values for δ13C and δ34S declined by -1.62 and -1.18‰ over the time of the study period, respectively, while values for δ15N showed no trend. Total Hg concentrations were positively related to both δ13C and δ34S. Yearly median THg concentrations ranged from 1.61 to 2.75 µg/g and increased nonlinearly by 0.86 µg/g in total over the study. Correcting THg concentrations for stable isotope values of carbon and sulfur and additionally breeding status and age slightly accelerated the increase in THg concentrations; however, confidence intervals of the raw THg trend and the corrected THg trend had substantial overlap. The rise in THg concentrations in the polar bear food web was possibly related to climate-related re-emissions of previously stored Hg from thawing sea-ice, glaciers, and permafrost.

The stress of Arctic warming on polar bears.

Arctic ecosystems are changing rapidly in response to climate warming. While Arctic mammals are highly evolved to these extreme environments, particularly with respect to their stress axis, some species may have limited capacity to adapt to this change. We examined changes in key components of the stress axis (cortisol and its carrier protein-corticosteroid binding globulin [CBG]) in polar bears (Ursus maritimus) from western Hudson Bay (N = 300) over a 33 year period (1983-2015) during which time the ice-free period was increasing. Changing sea ice phenology limits spring hunting opportunities and extends the period of onshore fasting. We assessed the response of polar bears to a standardized stressor (helicopter pursuit, darting, and immobilization) during their onshore fasting period (late summer-autumn) and quantified the serum levels of the maximum corticosteroid binding capacity (MCBC) of CBG, the serum protein that binds cortisol strongly, and free cortisol (FC). We quantified bear condition (age, sex, female with cubs or not, fat condition), sea ice (breakup in spring-summer, 1 year lagged freeze-up in autumn), and duration of fasting until sample collection as well as cumulative impacts of the latter environmental traits from the previous year. Data were separated into "good" years (1983-1990) when conditions were thought to be optimal and "poor" years (1991-2015) when sea ice conditions deteriorated and fasting on land was extended. MCBC explained 39.4% of the variation in the good years, but only 28.1% in the poor ones, using both biological and environmental variables. MCBC levels decreased with age. Changes in FC were complex, but more poorly explained. Counterintuitively, MCBC levels increased with increased time onshore, 1 year lag effects, and in poor ice years. We conclude that MCBC is a biomarker of stress in polar bears and that the changes we document are a consequence of climate warming.

Contrasting Temporal Patterns of Mercury, Niche Dynamics, and Body Fat Indices of Polar Bears and Ringed Seals in a Melting Icescape.

Polar bears (Ursus maritimus) and ringed seals (Pusa hispida) have a strong predator-prey relationship and are facing climate-associated Arctic habitat loss and harmful dietary exposure to total mercury (THg) and other pollutants. However, little is known about whether both species inhabiting the same area exhibit similar temporal patterns in Hg concentration, niche dynamics, and body fat indices. We used THg, δ13C, and δ15N values of western Hudson Bay polar bear hair (2004-2016) and ringed seal muscle samples (2003-2015) to investigate temporal trends of these variables and multidimensional niche metrics, as well as body fat indices for both species. We found a decline in THg concentration (by 3.8% per year) and δ13C (by 1.5‰) in ringed seals suggesting a change in feeding habits and carbon source use over time, whereas no significant changes occurred in polar bears. In contrast, the polar bear 3-dimensional niche size decreased by nearly half with no change in ringed seal niche size. The δ13C spacing between both species increased by approximately 1.5× suggesting different responses to annual changes in sympagic-pelagic carbon source production. Ringed seal body fat index was higher in years of earlier sea ice breakup with no change occurring in polar bears. These findings indicate that both species are responding differently to a changing environment suggesting a possible weakening of their predator-prey relationship in western Hudson Bay.

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.

The use of hair as a proxy for total and methylmercury burdens in polar bear muscle tissue.

Polar bears (Ursus maritimus) are an ecologically important species in the Arctic, whose health, and that of the people whose livelihoods depend on them, are increasingly affected by climate change and the bioaccumulation of contaminants such as mercury (Hg). Although methylmercury (MeHg) is the toxic form of Hg that biomagnifies up food webs, risk assessment studies typically only report on total Hg (THg) concentrations because it is cheaper to quantify. Furthermore, hair is commonly analysed for THg in polar bear as well as human risk assessment studies because it is relatively non-invasive to collect, yet we know little of how THg and MeHg concentrations differ between hair and muscle tissues. In this study, we quantified THg and MeHg concentrations in hair and muscle from 44 polar bears (24 sub-adults: 9 females, 15 males; 18 adults: 5 females, 13 males, and 2 males of unknown age group), harvested in 2015 and 2016 from four subpopulations in Nunavut, Canada (Davis Strait, n = 3; Gulf of Boothia, n = 8; Baffin Bay, n = 15; Foxe Basin, n = 18). We found only moderately positive correlations (0.4 ≤ r ≤ 0.5) between THg concentrations in hair and THg and MeHg concentrations in muscle. Further, 75% and 88% of THg was MeHg in hair and muscle, respectively. High concentrations of THg in hair - 71% of the samples were above the suggested neurochemical no observed effect level for polar bears - suggest some of the bears may be adversely affected by Hg-related health effects. Despite this, all MeHg concentrations in muscle (0.1 to 0.4 mg/kg (wet weight, ww)) were below the consumption maximum Hg concentration of 0.5 mg/kg (ww) set by Canadian health authorities.

Chronic stress and body condition of wolf-killed prey in Prince Albert National Park, Saskatchewan.

Chronic stress and poor body condition can cause adverse physiological and behavioural responses and may make animals more vulnerable to predation. We examined hair cortisol concentration (HCC) and marrow lipid content, as bioindicators of chronic stress and body condition, respectively, of bison (Bison bison bison), moose (Alces alces) and white-tailed deer (Odocoileus virginianus) killed by wolves (Canis lupus) in Prince Albert National Park (PANP), Saskatchewan, Canada. The Sturgeon River plains bison population in PANP is one of only a few wild populations of plains bison in their historical range in Canada and has experienced a decline of around 50% since 2005. We expected wolf-killed bison to have elevated HCC compared to human-harvested bison and that there would be a negative relationship between HCC and marrow lipids among wolf-killed animals. We compared HCC between different mortality sources for bison (wolf-killed n = 20 or human-harvested n = 23) and found that HCC was significantly elevated in wolf-killed bison (¯ = 7.56 ± 1.35 pg/mg). We found that HCC, species, sex and snow depth were all significant predictor variables of marrow lipid content of bison (n = 14), moose (n = 11) and deer (n = 27). Bison displayed the strongest negative correlation between HCC and marrow lipid content (r2 = 0.31). Our results suggest that chronic stress and poor body condition make prey more vulnerable to predation by wolves. HCC and marrow lipid content can provide reliable indicators of the physiological response of animals to stressors and may provide information on expected predator success that can be used to predict predator population dynamics.

How many cubs can a mum nurse? Maternal age and size influence litter size in polar bears.

Life-history theory predicts that females' age and size affect the level of maternal investment in current reproduction, balanced against the future reproductive effort, maintenance and survival. Using long-term (30 years) individual data on 193 female polar bears ( Ursus maritimus), we assessed age- and size-specific variation on litter size. Litter size varied with maternal age, younger females had higher chances of losing a cub during their first months of life. Results suggest an improvement in reproductive abilities early in life due to experience with subsequent reproductive senescence. Litter size increased with maternal size, indicating that size may reflect individual quality. We also found an optimum in the probability of having twins, suggesting stabilizing selection on female body size. Heterogeneity was observed among the largest females, suggesting that large size comes at a cost.

Modeling optimal responses and fitness consequences in a changing Arctic.

Animals must balance a series of costs and benefits while trying to maximize their fitness. For example, an individual may need to choose how much energy to allocate to reproduction versus growth, or how much time to spend on vigilance versus foraging. Their decisions depend on complex interactions between environmental conditions, behavioral plasticity, reproductive biology, and energetic demands. As animals respond to novel environmental conditions caused by climate change, the optimal decisions may shift. Stochastic dynamic programming provides a flexible modeling framework with which to explore these trade-offs, but this method has not yet been used to study possible changes in optimal trade-offs caused by climate change. We created a stochastic dynamic programming model capturing trade-off decisions required by an individual adult female polar bear (Ursus maritimus) as well as the fitness consequences of her decisions. We predicted optimal foraging decisions throughout her lifetime as well as the energetic thresholds below which it is optimal for her to abandon a reproductive attempt. To explore the effects of climate change, we shortened the spring feeding period by up to 3 weeks, which led to predictions of riskier foraging behavior and higher reproductive thresholds. The resulting changes in fitness may be interpreted as a best-case scenario, where bears adapt instantaneously and optimally to new environmental conditions. If the spring feeding period was reduced by 1 week, her expected fitness declined by 15%, and if reduced by 3 weeks, expected fitness declined by 68%. This demonstrates an effective way to explore a species' optimal response to a changing landscape of costs and benefits and highlights the fact that small annual effects can result in large cumulative changes in expected lifetime fitness.

Predation risk and space use of a declining Dall sheep (Ovis dalli dalli) population.

The abundance of ungulate populations may fluctuate in response to several limiting factors, including climate, diseases, and predation. In the northern Richardson Mountains, Canada, Dall sheep (Ovis dalli dalli) have undergone a major decline in the past decades and predation by grizzly bears (Ursus arctos) and wolves (Canis lupus) was suspected as a leading cause. To better understand the relationship between these three species located in this rugged and remote ecosystem, we relied on a combination of indirect methods. We investigated the apparent role of predation on the Dall sheep population using spatial ecology and stable isotopes. We examined seasonal variation in predation risk, focusing on how it may affect Dall sheep habitat use and sexual segregation, and we evaluated the proportion of Dall sheep in the diet of both predators using stable isotopes. The movements of the three species were monitored by satellite telemetry. Dall sheep habitat use patterns were analyzed using topographical features, greenness index, land cover, and apparent predation risk. The diets of grizzly bears and wolves were examined using a Bayesian mixing model for carbon and nitrogen stable isotopes. We found that Dall sheep habitat use varied seasonally, with different patterns for ewes and rams. Exposure to grizzly bear risk was higher for rams during summer, while ewes were further exposed to wolf apparent predation risk during winter. The importance of safe habitats for ewes was reflected in space use patterns. Stable isotopes analyses suggested that the diet of grizzly bears was largely from animal sources, with mountain mammals comprising about one quarter. Wolves mostly fed on both aquatic browsers and mountain mammals. Diet variation between individual predators suggested that some individuals specialized on mountain mammals, likely including Dall sheep. We conclude that grizzly bear and wolf apparent predation risk are important in driving Dall sheep habitat use and play a role in sexual segregation. Overall, this study presents an innovative combination of indirect methods that could be applied elsewhere to better understand predator-prey dynamics in remote ecosystems.

Ringed seal demography in a changing climate.

Climate change is affecting species' distributions and abundances worldwide. Baseline population estimates, against which future observations may be compared, are necessary if we are to detect ecological change. Arctic sea ice ecosystems are changing rapidly and we lack baseline population estimates for many ice-associated species. Provided we can detect them, changes in Arctic marine ecosystems may be signaled by changes in indicator species such as ringed seals (Pusa hispida). Ringed seal monitoring has provided estimates of survival and fertility rates, but these have not been used for population-level inference. Using matrix population models, we synthesized existing demographic parameters to obtain estimates of historical ringed seal population growth and structure in Amundsen Gulf and Prince Albert Sound, Canada. We then formalized existing hypotheses about the effects of emerging environmental stressors (i.e., earlier spring ice breakup and reduced snow depth) on ringed seal pup survival. Coupling the demographic model to ice and snow forecasts available from the Coupled Model Intercomparison Project resulted in projections of ringed seal population size and structure up to the year 2100. These projections showed median declines in population size ranging from 50% to 99%. Corresponding to these projected declines were substantial changes in population structure, with increasing proportions of ringed seal pups and adults and declining proportions of juveniles. We explored if currently collected, harvest-based data could be used to detect the projected changes in population stage structure. Our model suggests that at a present sample size of 100 seals per year, the projected changes in stage structure would only be reliably detected by mid-century, even for the most extreme climate models. This modeling process revealed inconsistencies in existing estimates of ringed seal demographic rates. Mathematical population models such as these can contribute both to understanding past population trends as well as predicting future ones, both of which are necessary if we are to detect and interpret future observations.