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USGS2022

Tooth wear and the apparent consumption of human foods among American black bears (Ursus americanus) in Great Smoky Mountains National Park, USA

Stable isotope analyses of hair have been used to estimate the consumption of human foods by American black bears ( Ursus americanus ). Consumption of human foods influences body mass and reproductive success of bears. However, the underlying factors that cause some bears to become conflict bears and resort to consuming human foods as a portion of their diet are not fully understood. We collected hair samples for stable isotope analysis from 51 black bears in Great Smoky Mountains National Park, Tennessee, USA in 2006. We used δ 13 C values of hairs to determine if the bears were consuming C 3 -based (natural foods) or C 4 -based (human foods) diets, and δ 15 N values, which increase with more meat in the diet, as a further indication of the consumption of human foods. Male bears with the heaviest tooth wear had a combination of higher δ 15 N and δ 13 C values, suggesting that they consume human foods to a greater extent than do other black bears. Based on our results, we hypothesize that tooth wear, and thus dental health, may play a role in the consumption of human foods by larger, male bears.

Tennessee↗

USGS2021

Incorporating climate change in a harvest risk assessment for polar bears Ursus maritimus in Southern Hudson Bay

Arctic marine mammals are harvested by Indigenous people for subsistence and are socially and culturally important. For ice-dependent species like the polar bear Ursus maritimus , management and conservation require understanding interactions between harvest and sea-ice loss due to climate change. We developed a demographic model to evaluate harvest risk for polar bears in Southern Hudson Bay, Canada, where the annual ice-free season has increased by approximately one month in recent decades. The model was based on the theta-logistic equation and allowed for density-dependent changes (through carrying capacity [ K ]) and density-independent changes (through population growth rate [ r ]). Model parameters were estimated using a Bayesian Monte Carlo method that included capture-recapture, aerial survey, and harvest data. Harvest management followed a state-dependent approach under which new estimates of abundance were used to update the harvest level every five years. Under a middle-of-the-road environmental scenario that assumed K and r would decline in proportion to projected sea-ice declines, annual removal of 0.02–0.03 of females resulted in a 0.8 probability of maintaining subpopulation abundance above maximum net productivity level for three polar bear generations (~34 years), our primary criterion for sustainability. Under more pessimistic and optimistic environmental scenarios, comparable female harvest rates were 0.01 and 0.055, respectively. Our coupled modeling-management framework can be used to inform tradeoffs between protection and sustainable use for wildlife populations experiencing habitat loss.

Southern Hudson Bay↗

USGS2006

Assessment of pesticide residues in army cutworm moths (Euxoa auxiliaris) from the Greater Yellowstone Ecosystem and their potential consequences to foraging grizzly bears (Ursus arctos horribilis)

During summer, a grizzly bear ( Ursus arctos horribilis ) in the Greater Yellowstone Ecosystem (GYE) (USA) can excavate and consume millions of army cutworm moths ( Euxoa auxiliaris ) (ACMs) that aggregate in high elevation talus. Grizzly bears in the GYE were listed as threatened by the US Fish and Wildlife Service in 1975 and were proposed for delisting in 2005. However, questions remain about key bear foods. For example, ACMs are agricultural pests and concern exists about whether they contain pesticides that could be toxic to bears. Consequently, we investigated whether ACMs contain and transport pesticides to bear foraging sites and, if so, whether these levels could be toxic to bears. In 1999 we collected and analyzed ACMs from six bear foraging sites. ACMs were screened for 32 pesticides with gas chromatography with electron capture detection (GC–ECD). Because gas chromatography with tandem mass spectrometry (GC–MS/MS) can be more sensitive than GC–ECD for certain pesticides, we revisited one site in 2001 and analyzed these ACMs with GC–MS/MS. ACMs contained trace or undetectable levels of pesticides in 1999 and 2001, respectively. Based on chemical levels in ACMs and numbers of ACMs a bear can consume, we calculated the potential of chemicals to reach physiological toxicity. These calculations indicate bears do not consume physiologically toxic levels of pesticides and allay concerns they are at risk from pesticides transported by ACMs. If chemical control of ACMs changes in the future, screening new ACM samples taken from bear foraging sites may be warranted.

Wyoming↗

USGS2020

Investigation of bed and den site selection by American black bears (Ursus americanus) in a landscape impacted by forest restoration treatments and wildfires

The combined effects of long-term fire suppression, logging, and overgrazing have negatively impacted many southwestern U.S. forests, resulting in decreased habitat quality for wildlife, and more frequent and severe wildfires. In response, land management agencies are implementing large-scale forest restoration treatments, but data on how wildlife respond to restoration treatments and wildfires are often limited. We investigated bed and den site selection of American black bears ( Ursus americanus ) using GPS location data and a use/available study design to assess the influence of habitat characteristics, including wildfires, prescribed burns, and thinning treatments on bed and den site selection in the Jemez Mountains, New Mexico. The most supported models suggested that black bears were more likely to select bed sites with a combination of low horizontal visibility ( β = −0.007, SE = 0.002; P = 0.002) and high stand basal area ( β = 0.013, SE = 0.005; P = 0.004). The highest-ranking model for den site selection indicated that black bears were more likely to select den sites with low horizontal visibility ( β = −0.0102, SE = 0.004; P = 0.006). Black bears used all disturbed sites to varying degrees (45% of study area), although 48% of bed sites were located in undisturbed habitat (55% of study area) while only 11% and 2% of bed sites were located in thinned and prescribed burn sites, respectively. Thirty-nine percent of bed sites were located in previous wildfire locations; however, 67% of these sites were in areas with low burn severity. Thirty-eight percent of den sites were located in previously disturbed habitat, 8 of these sites were burned by wildfires. In order to develop effective management plans for black bears, it is essential to understand responses to landscape-scale habitat disturbances due to wildfires and restoration activities, all of which are becoming more prevalent and widespread across southwestern forests. Accounting for the timing, size, and proximity of future restoration efforts would aid in mitigating potential short-term negative effects on black bears.

New Mexico↗

USGS2015

Establishing a definition of polar bear (Ursus maritimus) health: A guide to research and management activities

The meaning of health for wildlife and perspectives on how to assess and measure health, are not well characterized. For wildlife at risk, such as some polar bear (Ursus maritimus) subpopulations, establishing comprehensive monitoring programs that include health status is an emerging need. Environmental changes, especially loss of sea ice habitat, have raised concern about polar bear health. Effective and consistent monitoring of polar bear health requires an unambiguous definition of health. We used the Delphi method of soliciting and interpreting expert knowledge to propose a working definition of polar bear health and to identify current concerns regarding health, challenges in measuring health, and important metrics for monitoring health. The expert opinion elicited through the exercise agreed that polar bear health is defined by characteristics and knowledge at the individual, population, and ecosystem level. The most important threats identified were in decreasing order: climate change, increased nutritional stress, chronic physiological stress, harvest management, increased exposure to contaminants, increased frequency of human interaction, diseases and parasites, and increased exposure to competitors. Fifteen metrics were identified to monitor polar bear health. Of these, indicators of body condition, disease and parasite exposure, contaminant exposure, and reproductive success were ranked as most important. We suggest that a cumulative effects approach to research and monitoring will improve the ability to assess the biological, ecological, and social determinants of polar bear health and provide measurable objectives for conservation goals and priorities and to evaluate progress.

Science of the Total Environment↗

USGS2021

Measuring adrenal and reproductive hormones in hair from southern Beaufort Sea polar bears (Ursus maritimus)

Polar bears ( Ursus maritimus ) use sea ice to access marine mammal prey. In Alaska’s Southern Beaufort Sea, the declining availability of sea ice habitat in summer and fall has reduced opportunities for polar bears to routinely hunt on the ice for seals, their primary prey. This reduced access to prey may result in physiological stress with subsequent potential consequences to reproductive function (physiological changes that accompany reproduction), which can be measured via reproductive hormones. Hormone concentrations in hair can be used as a minimally invasive alternative to serum concentrations, which must come from animal captures. Hair samples also provide a long-term average measurement of hormone concentrations that is not influenced by short-term fluctuations like that of serum. The aim of this study was (1) to determine if a radioimmunoassay could be used to measure adrenal and reproductive hormones in polar bear hair, and (2) to determine what the relationship is between these hormones and other reproductive, condition, and demographic parameters of polar bears. We successfully validated this method for cortisol, progesterone, estradiol, and testosterone through the analysis of hair and serum of 141 free-ranging polar bears. We found that while hair cannot be used to estimate serum hormone concentrations during the breeding season, hormone concentrations in hair can be used to measure reproductive function in polar bears. Further, our findings support trends in previous studies measuring hormone concentrations in serum. We found that adrenal and some reproductive hormones were positively correlated in hair samples of females. Associations between hormone concentrations in hair and serum did not vary relative to reproductive status of adult females. Serum testosterone increased throughout the breeding season for adult males and was significantly associated with body mass index (BMI). Our research supports the use of hair as a measure of reproductive function in polar bears and allows us to monitor the future effects of climate change on polar bear physiology.

Alaska, Yukon↗

USGS1997

Use of ungulates by Yellowstone grizzly bears Ursus arctos

Previous results of fecal analysis from the Yellowstone area and the known abilities of grizzly bears Ursus arctos to acquire and digest tissue from vertebrates suggested that grizzlies in this ecosystem obtained substantial energy from ungulates. This issue was addressed using observations from radio-marked grizzly bears, 1977–1992. Ungulates potentially contributed the majority of energy required for activity during the non-denning season for both adult female and male grizzlies. Most of this energy (95%) was estimated to come from the largest-bodied ungulate species (elk Cervus elaphus , bison Bison bison , and moose Alces alces ), with greatest proportional contributions by scavenged adult male bison (16%), scavenged calf and yearling elk (10%) and adult female elk that were killed (8%) or scavenged (8%). Grizzlies acquired 30% of total edibles from ungulates by predation, of which 13% (or 4% of the total) came from predation on elk calves. Most scavenging occurred during the spring and was associated with the abundance and relative availability of different types of carrion. Predation and scavenging did not appear to be compensatory. Rather, total consumption of ungulates varied inversely with consumption of whitebark pine Pinus albicaulis seeds. The relative frequency of predation to scavenging increased with ungulate density. Contrary to previous suppositions, neither total ungulate use nor frequency of predation increased during the study, despite large increases in some ungulate populations. As expected by the identified trade-offs, Yellowstone grizzlies seemed to prey selectively upon moose, probably because of their solitary habits and forested surroundings, but otherwise favored vulnerable smaller-bodied ungulates such as elk calves. No predation on adult bison was observed.

Idaho, Montana, Wyoming↗

USGS1999

Defining landscapes suitable for restoration of grizzly bears Ursus arctos in Idaho

Informed management of large carnivores depends on the timely and useful presentation of relevant information. We describe an approach to evaluating carnivore habitat that uses pre-existing qualitative and quantitative information on humans and carnivores to generate coarse-scale maps of habitat suitability, habitat productivity, potential reserves, and areas of potential conflict. We use information pertinent to the contemplated reintroduction of grizzly bears Ursus arctos horribilis into central Idaho to demonstrate our approach. The approach uses measures of human numbers, their estimated distribution, road and trail access, and abundance and quality of bear foods to create standardized indices that are analogues of death and birth rates, respectively; the first subtracted from the second indicates habitat suitability ( HS ). We calibrate HS to sightings of grizzly bears in two ecosystems in northern Idaho and develop an empirical model from these same sightings based on piece-wise treatment of the variables contained in HS . Depending on whether the empirical model or HS is used, we estimate that there is 14 800 km 2 of suitable habitat in two blocks or 37 100 km 2 in one block in central Idaho, respectively. Both approaches show suitable habitat in the current Evaluation Area and in an area of southeastern Idaho centered on the Palisades Reservoir. Areas of highly productive habitat are concentrated in northern and western Idaho and in the Palisades area. Future conflicts between humans and bears are most likely to occur on the western and northern margins of suitable habitat in central Idaho, rather than to the east, where opposition to reintroduction of grizzly bears is currently strongest.

Idaho↗

USGS2003

Geographical distribution of organochlorine pesticides (OCPs) in polar bears (Ursus maritimus) in the Norwegian and Russian Arctic

Geographical variation of organochlorine pesticides (OCPs) was studied in blood samples from 90 adult female polar bear (Ursus maritimus) from Svalbard, Franz Josef Land, Kara Sea, East-Siberian Sea and Chukchi Sea. In all regions, oxychlordane was the dominant OCP. Regional differences in mean levels of HCB, oxychlordane, trans-nonachlor, ??-HCH, ??-HCH and p,p???-DDE were found. The highest levels of oxychlordane, trans-nonachlor and DDE were found in polar bears from Franz Josef Land and Kara Sea. HCB level was lowest in polar bears from Svalbard. Polar bears from Chukchi Sea had the highest level of ??- and ??-HCH. The lowest ??-HCH concentration was found in bears from Kara Sea. In all the bears, ???HCHs was dominated by ??-HCH. The geographical variation in OCP levels and pattern may suggest regional differences in pollution sources and different feeding habits in the different regions. Polar bears from the Western Russian Arctic were exposed to higher levels of chlordanes and p,p???-DDE than polar bears from locations westwards and eastwards from this region. This may imply the presence of a significant pollution source in the Russian Arctic area. The study suggests that the western Russian Arctic is the most contaminated region of the Arctic and warrants further research. ?? 2002 Elsevier Science B.V. All rights reserved.

Science of the Total Environment↗

USGS2003

The influence of diet on faecal DNA amplification and sex identification in brown bears ( Ursus arctos )

To evaluate the influence of diet on faecal DNA amplification, 11 captive brown bears (Ursus arctos) were placed on six restricted diets: grass (Trifolium spp., Haplopappus hirtus and Poa pratensis), alfalfa (Lupinus spp.), carrots (Daucus spp.), white-tailed deer (Odocoileus virginianus), blueberries (Vaccinium spp.) and salmon (Salmo spp.). DNA was extracted from 50 faecal samples of each restricted diet, and amplification of brown bear DNA was attempted for a mitochondrial DNA (mtDNA) locus and nuclear DNA (nDNA) locus. For mtDNA, no significant differences were observed in amplification success rates across diets. For nDNA, amplification success rates for salmon diet extracts were significantly lower than all other diet extracts (P < 0.001). To evaluate the accuracy of faecal DNA sex identification when female carnivores consume male mammalian prey, female bears were fed male white-tailed deer. Four of 10 extracts amplified, and all extracts were incorrectly scored as male due to amplification of X and Y-chromosome fragments. The potential biases highlighted in this study have broad implications for researchers using faecal DNA for individual and sex identification, and should be evaluated in other species.

Molecular Ecology↗

USGS2023

A body composition model with multiple storage compartments for polar bears (Ursus maritimus)

Climate warming is rapidly altering Arctic ecosystems. Polar bears ( Ursus maritimus ) need sea ice as a platform from which to hunt seals, but increased sea-ice loss is lengthening periods when bears are without access to primary hunting habitat. During periods of food scarcity, survival depends on the energy that a bear has stored in body reserves, termed storage energy, making this a key metric in predictive models assessing climate change impacts on polar bears. Here, we developed a body composition model for polar bears that estimates storage energy while accounting for changes in storage tissue composition. We used data of dissected polar bears ( n = 31) to link routinely collected field measures of total body mass and straight-line body length to the body composition of individual bears, described in terms of structural mass and two storage compartments, adipose and muscle. We then estimated the masses of metabolizable proteins and lipids within these storage compartments, giving total storage energy. We tested this multi-storage model by using it to predict changes in the lipid stores from an independent dataset of wild polar bears ( n = 36) that were recaptured 8–200 days later. Using length and mass measurements, our model successfully predicted direct measurements of lipid changes via isotopic dilutions (root mean squared error of 14.5 kg). Separating storage into two compartments, and allowing the molecular composition of storage to vary, provides new avenues for quantifying energy stores of individuals across their life cycle. The multi-storage body composition model thus provides a basis for further exploring energetic costs of physiological processes that contribute to individual survival and reproductive success. Given bioenergetic models are increasingly used as a tool to predict individual fitness and population dynamics, our approach for estimating individual energy stores could be applicable to a wide range of species.

Article↗

USGS2023

Serum virome of southern Beaufort Sea polar bears (Ursus maritimus) during a period of rapid climate change

Climate change affects the behavior, physiology and life history of many Arctic wildlife species. It can also influence the distribution and ecology of infectious agents. The southern Beaufort Sea (SB) subpopulation of polar bears ( Ursus maritimus ) has experienced dramatic behavioral changes due to retreating sea ice and other climate-related factors, but the effects of these changes on physiology and infection remain poorly understood. Using serum from polar bears sampled between 2004 and 2015 and metagenomic DNA sequencing, we identified 48 viruses, all of the family Anelloviridae . Anelloviruses are small, ubiquitous infectious agents with circular single-stranded DNA genomes that are not known to cause disease but, in humans, covary in diversity and load with immunological compromise. We therefore examined the usefulness of anelloviruses as biomarkers of polar bear physiological stress related to climate and habitat use. Polar bear anelloviruses sorted into two distinct clades on a phylogenetic tree, both of which also contained anelloviruses of giant pandas ( Ailuropoda melanoleuca ), another ursid. Neither anellovirus diversity nor load were associated with any demographic variables, behavioral factors or direct physiological measures. However, pairwise genetic distances between anelloviruses were positively correlated with pairwise differences in sampling date, suggesting that the polar bear “anellome”is evolving over time. These findings suggest that anelloviruses are not a sensitive indicator of polar physiological stress, but they do provide a baseline for evaluating future changes to polar bear viromes.

Alaska↗

USGS2023

Long-term assessment of relationships between changing environmental conditions and the physiology of southern Beaufort Sea polar bears (Ursus maritimus)

Climate change is influencing polar bear ( Ursus maritimus ) habitat, diet, and behavior but the effects of these changes on their physiology is not well understood. Blood-based biomarkers are used to assess the physiologic health of individuals but their usefulness for evaluating population health, especially as it relates to changing environmental conditions, has rarely been explored. We describe links between environmental conditions and physiologic functions of southern Beaufort Sea polar bears using data from blood samples collected from 1984 to 2018, a period marked by extensive environmental change. We evaluated associations between 13 physiologic biomarkers and circumpolar (Arctic oscillation index) and regional (wind patterns and ice-free days) environmental metrics and seasonal and demographic co-variates (age, sex, season, and year) known to affect polar bear ecology. We observed signs of dysregulation of water balance in polar bears following years with a lower annual Arctic oscillation index. In addition, liver enzyme values increased over time, which is suggestive of potential hepatocyte damage as the Arctic has warmed. Biomarkers of immune function increased with regional-scale wind patterns and the number of ice-free days over the Beaufort Sea continental shelf and were lower in years with a lower winter Arctic oscillation index, suggesting an increased allocation of energetic resources for immune processes under these conditions. We propose that the variation in polar bear immune and metabolic function is likely indicative of physiologic plasticity, a response that allows polar bears to remain in homeostasis even as they experience changes in nutrition and habitat in response to changing environments.

Alaska↗

USGS2009

Tetranucleotide microsatellite loci from the black bear (Ursus americanus)

We describe primers and polymerase chain reaction conditions to amplify 21 tetranucleotide microsatellite DNA loci in black bears (Ursus americanus). We tested primers using individuals from two populations, one each in Georgia and Florida. Among individuals from Georgia (n = 29), primer pairs yielded an average of 2.9 alleles (range, one to four) and an average observed heterozygosity (HO) of 0.50 (range, 0.00 to 0.79). Among individuals from Florida (n = 19), primer pairs yielded an average of 5.7 alleles (range, one to 14) and an HO of 0.55 (range, 0.00 to 1.00). A comparison of previously developed markers with individuals from Georgia suggests that bear populations in Georgia and Florida have reduced allelic diversity relative to other populations. ?? 2008 The Authors.

Molecular Ecology Resources↗

USGS2025

The American black bear (Ursus americanus) as an apex predator: Investigating the ecological role of the world’s most abundant large carnivore

Introduction American black bears ( Ursus americanus ) have been documented to have top-down effects, both consumptive (through predation) and nonconsumptive (through fear effects). However, their behavioural and dietary adaptability has led to uncertainty about the conditions under which these may occur. Objectives We aimed to (1) investigate when, where, and how often black bears influence lower trophic levels, and (2) critically assess the experimental design of such studies to determine trends and biases. Methods We conducted a systematic literature search and used linear mixed-effects models to assess the relationships among prey age class, the presence of other large carnivores, and the percentage of ungulate mortality due to black bears. We also summarised the measured effects of black bears on other carnivores and the results of any experimental studies. Results We found a strong negative relationship between puma presence and the percentage of ungulate mortality due to black bears, but only in areas with at least one other large carnivore (e.g., wolves). Among 16 studies of black bears' impacts on other carnivores, most documented negative behavioural effects. While only 5% of studies had strong experimental design, bear removal consistently increased juvenile ungulate survival. Discussion and Synthesis Our findings highlight black bears' strong top-down effects, resembling or exceeding those of other large carnivores. These effects were greatest when no other large carnivores were present, but as highly effective kleptoparasites, black bears can alter the feeding habits of other large carnivores. Thus, we argue that ‘apex’ is a conditional state, not a species-wide status.

Mammal Review↗

USGS2014

Influence of drift and admixture on population structure of American black bears ( Ursus americanus ) in the Central Interior Highlands, USA, 50 years after translocation

Bottlenecks, founder events, and genetic drift often result in decreased genetic diversity and increased population differentiation. These events may follow abundance declines due to natural or anthropogenic perturbations, where translocations may be an effective conservation strategy to increase population size. American black bears ( Ursus americanus ) were nearly extirpated from the Central Interior Highlands, USA by 1920. In an effort to restore bears, 254 individuals were translocated from Minnesota, USA, and Manitoba, Canada, into the Ouachita and Ozark Mountains from 1958 to 1968. Using 15 microsatellites and mitochondrial haplotypes, we observed contemporary genetic diversity and differentiation between the source and supplemented populations. We inferred four genetic clusters: Source, Ouachitas, Ozarks, and a cluster in Missouri where no individuals were translocated. Coalescent models using approximate Bayesian computation identified an admixture model as having the highest posterior probability (0.942) over models where the translocation was unsuccessful or acted as a founder event. Nuclear genetic diversity was highest in the source (A R = 9.11) and significantly lower in the translocated populations (A R = 7.07-7.34; P = 0.004). The Missouri cluster had the lowest genetic diversity (A R = 5.48) and served as a natural experiment showing the utility of translocations to increase genetic diversity following demographic bottlenecks. Differentiation was greater between the two admixed populations than either compared to the source, suggesting that genetic drift acted strongly over the eight generations since the translocation. The Ouachitas and Missouri were previously hypothesized to be remnant lineages. We observed a pretranslocation remnant signature in Missouri but not in the Ouachitas.

Molecular Ecology↗

USGS2008

Genetic characterization of Kenai brown bears (Ursus arctos): Microsatellite and mitochondrial DNA control region variation in brown bears of the Kenai Peninsula, south central Alaska

We collected data from 20 biparentally inherited microsatellite loci, and nucleotide sequence from the maternally inherited mitochondrial DNA (mtDNA) control region, to determine levels of genetic variation of the brown bears (Ursus arctos L., 1758) of the Kenai Peninsula, south central Alaska. Nuclear genetic variation was similar to that observed in other Alaskan peninsular populations. We detected no significant inbreeding and found no evidence of population substructuring on the Kenai Peninsula. We observed a genetic signature of a bottleneck under the infinite alleles model (IAM), but not under the stepwise mutation model (SMM) or the two-phase model (TPM) of microsatellite mutation. Kenai brown bears have lower levels of mtDNA haplotypic diversity relative to most other brown bear populations in Alaska. ?? 2008 NRC.

Canadian Journal of Zoology↗

USGS2012

Long-distance swimming by polar bears (Ursus maritimus) of the southern Beaufort Sea during years of extensive open water

Polar bears ( Ursus maritimus Phipps, 1774) depend on sea ice for catching marine mammal prey. Recent sea-ice declines have been linked to reductions in body condition, survival, and population size. Reduced foraging opportunity is hypothesized to be the primary cause of sea-ice-linked declines, but the costs of travel through a deteriorated sea-ice environment also may be a factor. We used movement data from 52 adult female polar bears wearing Global Positioning System (GPS) collars, including some with dependent young, to document long-distance swimming (>50 km) by polar bears in the southern Beaufort and Chukchi seas. During 6 years (2004-2009), we identified 50 long-distance swims by 20 bears. Swim duration and distance ranged from 0.7 to 9.7 days (mean = 3.4 days) and 53.7 to 687.1 km (mean = 154.2 km), respectively. Frequency of swimming appeared to increase over the course of the study. We show that adult female polar bears and their cubs are capable of swimming long distances during periods when extensive areas of open water are present. However, long-distance swimming appears to have higher energetic demands than moving over sea ice. Our observations suggest long-distance swimming is a behavioral response to declining summer sea-ice conditions.

Canadian Journal of Zoology↗