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.

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.

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.

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.

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.

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.

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.

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.

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.

Temporal Trends of Persistent Organic Pollutants in Barents Sea Polar Bears ( Ursus maritimus) in Relation to Changes in Feeding Habits and Body Condition.

Temporal trends of persistent organic pollutants (POPs: PCBs, OH-PCBs, p, p'-DDE, HCB, ß-HCH, oxychlordane, BDE-47, and 153) in relation to changes in feeding habits and body condition in adult female polar bears ( Ursus maritimus) from the Barents Sea subpopulation were examined over 20 years (1997-2017). All 306 samples were collected in the spring (April). Both stable isotope values of nitrogen (δ15N) and carbon (δ13C) from red blood cells declined over time, with a steeper trend for δ13C between 2012 and 2017, indicating a decreasing intake of marine and high trophic level prey items. Body condition, based on morphometric measurements, had a nonsignificant decreasing tendency between 1997 and 2005, and increased significantly between 2005 and 2017. Plasma concentrations of BDE-153 and ß-HCH did not significantly change over time, whereas concentrations of Σ4PCB, Σ5OH-PCB, BDE-47, and oxychlordane declined linearly. Concentrations of p, p'-DDE and HCB, however, declined until 2012 and 2009, respectively, and increased thereafter. Changes in feeding habits and body condition did not significantly affect POP trends. The study indicates that changes in diet and body condition were not the primary driver of POPs in polar bears, but were controlled in large part by primary and/or secondary emissions of POPs.

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.

Choose Your Poison-Space-Use Strategy Influences Pollutant Exposure in Barents Sea Polar Bears.

Variation in space-use is common within mammal populations. In polar bears, Ursus maritimus, some individuals follow the sea ice (offshore bears) whereas others remain nearshore yearlong (coastal bears). We studied pollutant exposure in relation to space-use patterns (offshore vs coastal) in adult female polar bears from the Barents Sea equipped with satellite collars (2000-2014, n = 152). First, we examined the differences in home range (HR) size and position, body condition, and diet proxies (nitrogen and carbon stable isotopes, n = 116) between offshore and coastal space-use. Second, we investigated how HR, space-use, body condition, and diet were related to plasma concentrations of polychlorinated biphenyls (PCBs), organochlorine pesticides (OCPs) ( n = 113), perfluoroalkyl substances (PFASs; n = 92), and hydroxylated-PCBs ( n = 109). Offshore females were in better condition and had a more specialized diet than did coastal females. PCBs, OCPs, and hydroxylated-PCB concentrations were not related to space-use strategy, yet PCB concentrations increased with increasing latitude, and hydroxylated-PCB concentrations were positively related to HR size. PFAS concentrations were 30-35% higher in offshore bears compared to coastal bears and also increased eastward. On the basis of the results we conclude that space-use of Barents Sea female polar bears influences their pollutant exposure, in particular plasma concentrations of PFAS.

Hundreds of Unrecognized Halogenated Contaminants Discovered in Polar Bear Serum.

Exposure of polar bears (Ursus maritimus) to persistent organic pollutants was discovered in the 1970s, but recent evidence suggests the presence of unknown toxic chemicals in their blood. Protein and phospholipid depleted serum was stirred with polyethersulfone capillaries to extract a broad range of analytes, and nontarget mass spectrometry with "fragmentation flagging" was used for detection. Hundreds of analytes were discovered belonging to 13 classes, including novel polychlorinated biphenyl (PCB) metabolites and many fluorinated or chlorinated substances not previously detected. All analytes were detected in the oldest (mid-1980s) archived polar bear serum from Hudson Bay and Beaufort Sea, and all fluorinated classes showed increasing trends. A mouse experiment confirmed the novel PCB metabolites, suggesting that these could be widespread in mammals. Historical exposure and toxic risk has been underestimated, and these halogenated contaminants pose uncertain risks to this threatened species.

Evaluating random search strategies in three mammals from distinct feeding guilds.

Searching allows animals to find food, mates, shelter and other resources essential for survival and reproduction and is thus among the most important activities performed by animals. Theory predicts that animals will use random search strategies in highly variable and unpredictable environments. Two prominent models have been suggested for animals searching in sparse and heterogeneous environments: (i) the Lévy walk and (ii) the composite correlated random walk (CCRW) and its associated area-restricted search behaviour. Until recently, it was difficult to differentiate between the movement patterns of these two strategies. Using a new method that assesses whether movement patterns are consistent with these two strategies and two other common random search strategies, we investigated the movement behaviour of three species inhabiting sparse northern environments: woodland caribou (Rangifer tarandus caribou), barren-ground grizzly bear (Ursus arctos) and polar bear (Ursus maritimus). These three species vary widely in their diets and thus allow us to contrast the movement patterns of animals from different feeding guilds. Our results showed that although more traditional methods would have found evidence for the Lévy walk for some individuals, a comparison of the Lévy walk to CCRWs showed stronger support for the latter. While a CCRW was the best model for most individuals, there was a range of support for its absolute fit. A CCRW was sufficient to explain the movement of nearly half of herbivorous caribou and a quarter of omnivorous grizzly bears, but was insufficient to explain the movement of all carnivorous polar bears. Strong evidence for CCRW movement patterns suggests that many individuals may use a multiphasic movement strategy rather than one-behaviour strategies such as the Lévy walk. The fact that the best model was insufficient to describe the movement paths of many individuals suggests that some animals living in sparse environments may use strategies that are more complicated than those described by the standard random search models. Thus, our results indicate a need to develop movement models that incorporate factors such as the perceptual and cognitive capacities of animals.

Hair Mercury Concentrations in Western Hudson Bay Polar Bear Family Groups.

Methylmercury is one of the more toxic forms of mercury (Hg), the biomagnification of which is prevalent in the Arctic where apex predators such as polar bears (Ursus maritimus) can carry high loads. The maternal transfer of contaminants to offspring is a concern, as offspring may be particularly sensitive to the effects of environmental pollutants during early development. However, few studies of polar bears report on Hg in dependent young. We examined hair total Hg (THg) concentrations in 24 polar bear family groups in western Hudson Bay: mother, cub-of-the-year (COY), yearling, and 2 year old. THg concentrations increased with bear age, with COYs having lower concentrations than other offspring groups (p ≤ 0.008). Using AICc-based regression models, we found maternal THg to be positively related to body condition and litter size, while overall offspring THg was positively related to maternal body condition in addition to being dependent on the sex and age of the offspring. COY THg concentrations were positively related to maternal THg while also depending on the sex of the offspring. Considering our results, future studies in polar bear ecotoxicology are encouraged to include offspring of different ages and sexes.

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.

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.

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.

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.