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Todd C. Atwood

Publications and source records attributed to Todd C. Atwood.

At least 37 records · Page 2Linked to original sources

Fecal DNA metabarcoding shows credible short-term prey detections and explains variation in the gut microbiome of two polar bear subpopulations

This study developed and evaluated DNA metabarcoding to identify the presence of pinniped and cetacean prey DNA in fecal samples of East Greenland (EG) and Southern Beaufort Sea (SB) polar bears Ursus maritimus sampled in the spring of 2015-2019. Prey DNA was detected in half (49/92) of all samples, and when detected, ringed seal Pusa hispida was the predominant prey species, identified in 100% (22/22) of EG and 81% (22/27) of SB polar bear samples with prey DNA detected. Bearded seal Erignathus barbatus DNA was found in 19% (5/27) of SB polar bear samples for which prey DNA was detected. Prey DNA detection frequencies and relative abundances were compared to estimates from quantitative fatty acid signature analysis (QFASA) for a subset of SB polar bears. Ringed seal and bearded seal were the main prey identified by both methods, but QFASA also identified 2 cetacean prey species not found by prey DNA. Differences in DNA metabarcoding vs. QFASA results were likely related to the different dietary timescales captured by each approach, i.e. short-term vs. long-term diet, respectively. Prey DNA detection, sex/age class, and subpopulation significantly explained variation in polar bear gut bacterial composition. Polar bear samples with prey DNA detected were associated with higher abundances of the bacterial classes Clostridia and Bacilli and lower abundances of Negativicutes. Fecal DNA metabarcoding is thus useful for identifying recent prey of polar bears, complementing quantitative and likely longer-term QFASA estimates, and may help understand variation in the polar bear gut microbiome.

Marine Ecology Progress Series (MEPS)

Feasibility of implementing a long-term plan to monitor the Arctic Basin polar bear subpopulation

The Arctic Basin (AB) polar bear subpopulation is the least studied of the 19 global polar bear subpopulations. The Polar Bear Specialist Group (PBSG) recognizes the AB subpopulation as a regional grouping intended to include bears that do not belong to any of the remaining subpopulations that have data to support boundary delineations. Very little is currently known about the AB subpopulation including information on its size, genetic uniqueness, or movement patterns, most likely due to its remote range, and uncertainty about whether the AB subpopulation represents a true subpopulation. We provide an overview on the minimum requirements needed to monitor the AB subpopulation; challenges for establishing and implementing a monitoring program; feasibility of different monitoring techniques; alternative sources of inference; and general suggestions for moving forward.

Report

Efficacy of bear spray as a deterrent against polar bears

Although there have been few attempts to systematically analyze information on the use of deterrents on polar bears ( Ursus maritimus ), understanding their effectiveness in mitigating human-polar bear conflicts is critical to ensuring both human safety and polar bear conservation. To fill this knowledge gap, we analyzed 19 incidents involving the use of bear spray on free-ranging polar bears from 1986 to 2019 in Canada, Russia, and the United States to evaluate the effectiveness of bear spray as a polar bear deterrent. We found that bear spray was an effective deterrent in close-range encounters with polar bears, stopping undesirable behavior in 18 of 19 incidents. Bear spray effectively deterred both curious and aggressive polar bears, including polar bears attempting to attack people. The mean distance between user and bear at the time of spraying was 2 m (min–max = 0.2–10.0 m, mode = 1 m), though bears were usually first seen at greater distances. Bear spray was successfully deployed against polar bears in all 4 seasons. Wind affected spray performance in 1 of 19 of incidents. In 8 of 14 bear spray incidents, other deterrents were used without success before bear spray was used effectively to deter polar bears. No humans or polar bears were killed or injured in any of the incidents in which bear spray was used. We also analyzed 54 polar bear attacks and attempted attacks on humans where bear spray was not carried. The data suggest that in 93% of those incidents, the use of bear spray might have saved the lives of both the people and bears involved if it had been available and used. Our analysis improves our understanding of the effectiveness of bear spray for polar bear conflict mitigation.

Wildlife Society Bulletin

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 their energy needs. Here, we assessed whether polar bears maintained dietary energy density by prey switching in response to spatial-temporal 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.

Ecological Applications

Pleistocene–Holocene vicariance, not Anthropocene landscape change, explains the genetic structure of American black bear (Ursus americanus) populations in the American Southwest and northern Mexico

The phylogeography of the American black bear ( Ursus americanus ) is characterized by isolation into glacial refugia, followed by population expansion and genetic admixture. Anthropogenic activities, including overharvest, habitat loss, and transportation infrastructure, have also influenced their landscape genetic structure. We describe the genetic structure of the American black bear in the American Southwest and northern Mexico and investigate how prehistoric and contemporary forces shaped genetic structure and influenced gene flow. Using a suite of microsatellites and a sample of 550 bears, we identified 14 subpopulations organized hierarchically following the distribution of ecoregions and mountain ranges containing black bear habitat. The pattern of subdivision we observed is more likely a product of postglacial habitat fragmentation during the Pleistocene and Holocene, rather than a consequence of contemporary anthropogenic barriers to movement during the Anthropocene. We used linear mixed-effects models to quantify the relationship between landscape resistance and genetic distance among individuals, which indicated that both isolation by resistance and geographic distance govern gene flow. Gene flow was highest among subpopulations occupying large tracts of contiguous habitat, was reduced among subpopulations in the Madrean Sky Island Archipelago, where montane habitat exists within a lowland matrix of arid lands, and was essentially nonexistent between two isolated subpopulations. We found significant asymmetric gene flow supporting the hypothesis that bears expanded northward from a Pleistocene refugium located in the American Southwest and northern Mexico and that major highways were not yet affecting gene flow. The potential vulnerability of the species to climate change, transportation infrastructure, and the US–Mexico border wall highlights conservation challenges and opportunities for binational collaboration.

Arizona, New Mexico, Utah, Wyoming

Modeling the spatial and temporal dynamics of land-based polar bear denning in Alaska

Although polar bears ( Ursus maritimus ) of the Southern Beaufort Sea (SBS) subpopulation have commonly created maternal dens on sea ice in the past, maternal dens on land have become increasingly prevalent as sea ice declines. This trend creates conditions for increased human–bear interactions associated with local communities and industrial activity. Maternal denning is a vulnerable period in the polar bear life cycle, and den disturbance could lead to den abandonment, cub mortality, and negative population impacts. We used published long-term data to parameterize a Bayesian hierarchical model of annual land den abundance during 2000–2015, in 4 regions of northern Alaska, USA, with current or potential future oil and gas activity. We also estimated long-term (1982–2015) shifts in the spatial distribution of land dens within and among regions using kernel density estimation and assessed the influence of local and regional sea ice and snow conditions on den site selection using a random forest resource selection function. Our objectives were to quantify current den distribution and abundance, test for distributional shifts over time, and investigate if those shifts could be attributed to environmental variables related to den habitat. We estimated that between 2000 and 2015, the SBS contained a median 123 dens in a typical year, of which 68 occurred on land. The region between the Colville and Canning rivers, where most current oil and gas activity occurred, also contained the largest fraction of land dens. Overall, land dens were disproportionately concentrated on barrier islands and on land within 30 km of the coast. The probability of dens occurring on land varied from 1982–1999 to 2000–2015 in all regions, and the overall distribution of land dens shifted west between those periods. This regional-scale change in den distribution was predictable based on spatial and temporal heterogeneity in snow and sea ice conditions within 50 km of individual den locations. Land denning is likely to become increasingly common with continued sea ice loss, and our results and modeling framework could be used to design additional mitigation strategies for reducing the risk of incidental take due to den disturbance.

Alaska

Comparisons of Coupled Model Intercomparison Project Phase 5 (CMIP5) and Coupled Model Intercomparison Project Phase 6 (CMIP6) sea-ice projections in polar bear (Ursus maritimus) ecoregions during the 21st century

Climate model projections are commonly used to assess potential impacts of global warming on a breadth of social, economic, and environmental topics. Modeling centers throughout the world coordinate to apply a consistent suite of radiative forcing experiments so that all model outputs can be collectively analyzed and compared. Three generations of model outputs have been produced and made available to the scientific community through the Coupled Model Intercomparison Project (CMIP): CMIP3 disseminated during the mid-2000s, CMIP5 during the early-2010s, and CMIP6 during the late-2010s. Twenty-first century sea-ice projections from CMIP3 and CMIP5 models have been used in Bayesian network assessments of how climate change could impact the future persistence of polar bears ( Ursus maritimus ) throughout their range. In this report, we compare sea-ice projections by CMIP6 models to those of CMIP5 models in each of four polar bear ecoregions over the 21st century. We evaluate differences between the two CMIP generations with respect to other sources of variability that affect uncertainties of the model projections: (1) variability from different models; (2) variability from different greenhouse gas emissions scenarios; and (3) natural (internal) variability in the earth’s climate system. We found that natural variability as well as that attributable to models dominated uncertainties in sea-ice projections in all months and ecoregions during the first half of the 21st century, while emissions scenarios dominated uncertainties during the late 21st century. By comparison, we found only slight differences between the CMIP6 and CMIP5 model projections of sea ice. Applying CMIP6 instead of CMIP5 sea-ice projections to the polar bear Bayesian network model developed in 2016, therefore, would not qualitatively change the population status outcomes published therein.

Open-File Report

Evaluating the efficacy of aerial infrared surveys to detect artificial polar bear dens

The need to balance economic development with impacts to Arctic wildlife has been a prominent subject since petroleum exploration began on the North Slope of Alaska, USA, in the late 1950s. The North Slope region includes polar bears ( Ursus maritimus ) of the southern Beaufort Sea subpopulation, which has experienced a long-term decline in abundance. Pregnant polar bears dig dens in snow drifts during winter and are vulnerable to disturbance, as den abandonment and mortality of neonates may result. Maternal denning coincides with the peak season of petroleum exploration and construction, raising concerns that human activities may disrupt denning. To minimize disturbance of denning polar bears, aerial infrared (AIR) surveys are routinely used to search for dens within planned industry activity areas and that information is used to implement mitigation. Aerial infrared surveys target the heat signature emanating from dens. Despite use by industry for >15 years, the efficacy of AIR and the factors that impact its ability to detect dens remains uncertain. Here, we evaluate AIR using artificial dens and observers naïve to locations to estimate detection probability and its relationship with covariates including weather variables, den characteristics, infrared sensor and altitude, and survey order to identify potential evidence of in-flight observer learning occurring between surveys. In December 2019 we constructed 14 dens (each with an artificial heat source), and 11 control sites (disturbed sites without dens). Between December 2019 and January 2020, 3 survey crews flew 6 independent AIR surveys within the vicinity of dens and control sites and video-recorded AIR imagery. Observers identified putative dens either in flight or during post-flight review of recordings. We assessed detection probability with a simple Bayesian model using 3 subsets of data: 1) all detection/non-detection data; 2) detection/non-detection data restricted to instances where sample sites were confirmed to have been properly scanned by AIR during post-study verification (i.e., when den locations were known); and 3) all dens visible on the recorded imagery during post-study verification, even if they were not seen during the survey or during post-flight review. Subsets 1 and 2 most closely resembled den surveys flown for oil and gas industry and had detection probabilities of 0.15 (95% CI = 0.08–0.23) and 0.24 (95% CI = 0.13–0.37), respectively. Detection probability was 0.41 (95% CI = 0.25–0.58) for subset 3. Higher wind speeds and larger den volume negatively influenced detection probability. Our low detection rate compared to previous studies could partially be the result of differences in study design, such as survey flight patterns. Our results suggest that AIR, as it is currently used, is unlikely to detect most polar bear dens in surveyed areas. Resource managers who use AIR should consider a suite of additional methods (e.g., habitat mapping, probabilistic den distribution, AIR methodology improvements) for minimizing impacts of industry on denning polar bears.

Alaska

Marine mammal hotspots across the circumpolar Arctic

Aim Identify hotspots and areas of high species richness for Arctic marine mammals. Location Circumpolar Arctic. Methods A total of 2115 biologging devices were deployed on marine mammals from 13 species in the Arctic from 2005 to 2019. Getis-Ord G i * hotspots were calculated based on the number of individuals in grid cells for each species and for phylogenetic groups (nine pinnipeds, three cetaceans, all species) and areas with high species richness were identified for summer (Jun-Nov), winter (Dec-May) and the entire year. Seasonal habitat differences among species’ hotspots were investigated using Principal Component Analysis. Results Hotspots and areas with high species richness occurred within the Arctic continental-shelf seas and within the marginal ice zone, particularly in the “Arctic gateways” of the north Atlantic and Pacific oceans. Summer hotspots were generally found further north than winter hotspots, but there were exceptions to this pattern, including bowhead whales in the Greenland-Barents Seas and species with coastal distributions in Svalbard, Norway and East Greenland. Areas with high species richness generally overlapped high-density hotspots. Large regional and seasonal differences in habitat features of hotspots were found among species but also within species from different regions. Gap analysis (discrepancy between hotspots and IUCN ranges) identified species and regions where more research is required. Main conclusions This study identified important areas (and habitat types) for Arctic marine mammals using available biotelemetry data. The results herein serve as a benchmark to measure future distributional shifts. Expanded monitoring and telemetry studies are needed on Arctic species to understand the impacts of climate change and concomitant ecosystem changes (synergistic effects of multiple stressors). While efforts should be made to fill knowledge gaps, including regional gaps and more complete sex and age coverage, hotspots identified herein can inform management efforts to mitigate the impacts of human activities and ecological changes, including creation of protected areas.

Diversity and Distributions

The role of satellite telemetry data in 21st century conservation of polar bears (Ursus maritimus)

Satellite telemetry (ST) has played a critical role in the management and conservation of polar bears ( Ursus maritimus ) over the last 50 years. ST data provide biological information relevant to subpopulation delineation, movements, habitat use, maternal denning, health, human-bear interactions, and accurate estimates of vital rates and abundance. Given that polar bears are distributed at low densities over vast and remote habitats, much of the information provided by ST data cannot be collected by other means. Obtaining ST data for polar bears requires chemical immobilization and application of a tracking device. Although immobilization has not been found to have negative effects beyond a several-day reduction in activity, over the last few decades opposition to immobilization and deployment of satellite-linked radio collars has resulted in a lack of current ST data in many of the 19 recognized polar bear subpopulations. Here, we review the uses of ST data for polar bears and evaluate its role in addressing 21 st century conservation and management challenges, which include estimation of sustainable harvest rates, understanding the impacts of climate warming, delineating critical habitat, and assessing potential anthropogenic impacts from tourism, resource development and extraction. We found that in subpopulations where ST data have been consistently collected, information was available to estimate vital rates and subpopulation density, document the effects of sea-ice loss, and inform management related to subsistence harvest and regulatory requirements. In contrast, a lack of ST data in some subpopulations resulted in increased bias and uncertainty in ecological and demographic parameters, which has a range of negative consequences. As sea-ice loss due to climate warming continues, there is a greater need to monitor polar bear distribution, habitat use, abundance, and subpopulation connectivity. We conclude that continued collection of ST data will be critically important for polar bear management and conservation in the 21 st century and that the benefits of immobilizing small numbers of individual polar bears in order to deploy ST devices significantly outweigh the risks.

Frontiers in Marine Science

Distinct gut microbiomes in two polar bear subpopulations inhabiting different sea ice ecoregions

Gut microbiomes were analyzed by 16S rRNA gene metabarcoding for polar bears ( Ursus maritimus ) from the southern Beaufort Sea (SB), where sea ice loss has led to increased use of land-based food resources by bears, and from East Greenland (EG), where persistent sea ice has allowed hunting of ice-associated prey nearly year-round. SB polar bears showed a higher number of total (940 vs. 742) and unique (387 vs. 189) amplicon sequence variants and higher inter-individual variation compared to EG polar bears. Gut microbiome composition differed significantly between the two subpopulations and among sex/age classes, likely driven by diet variation and ontogenetic shifts in the gut microbiome. Dietary tracer analysis using fatty acid signatures for SB polar bears showed that diet explained more intrapopulation variation in gut microbiome composition and diversity than other tested variables, i.e., sex/age class, body condition, and capture year. Substantial differences in the SB gut microbiome relative to EG polar bears, and associations between SB gut microbiome and diet, suggest that the shifting foraging habits of SB polar bears tied to sea ice loss may be altering their gut microbiome, with potential consequences for nutrition and physiology.

Alaska

Observed and forecasted changes in land use by polar bears in the Beaufort and Chukchi Seas, 1985–2040

Monitoring changes in the distribution of large carnivores is important for managing human safety and supporting conservation. Throughout much of their range, polar bears ( Ursus maritimus ) are increasingly using terrestrial habitats in response to Arctic sea ice decline. Their increased presence in coastal areas has implications for bear-human conflict, inter-species interactions, and polar bear health and survival. We examined observed trends in land use over three decades by polar bears in the southern Beaufort Sea (SB) and Chukchi Sea (CS) where bears have traditionally spent most of the year on the sea ice. Using data from 408 adult females fitted with satellite radio-collars, we examined trends in the annual proportion of bears coming onshore (hereafter referred to as “percent of bears”) during the summer for ≥21 days, arrival and departure dates, duration spent onshore and relationships with sea ice metrics. We then estimated future land use through 2040 by extrapolating trends and by combining observed relationships between land use and sea ice with projections of future sea ice from an ensemble of earth system models. The observed percent of bears summering onshore and their duration onshore was correlated with the percent of open water that occurred within their population’s range between July and October. As sea ice declined, the percent of bears summering onshore increased from ~5 to 30% in the SB and ~10 to 50% in the CS and duration onshore increased by >30 days to 60–70 days in both populations. Using a range of greenhouse gas emission scenarios and adjustments for faster than forecasted sea ice loss we estimated that 50-62% of SB and 79-88% of CS bears will spend 90–108 and 110–126 days onshore during summer in the SB and CS, respectively, by 2040. Sea ice projections varied little between greenhouse gas emission scenarios prior to 2040 but diverged thereafter. Observed and forecasted increases in polar bear land occupancy puts more bears in proximity to human activities and settlements for longer durations while extending the lack of access to their primary prey. Because human conflict is one of the primary factors affecting the conservation of large carnivores worldwide, mitigation of bear-human interactions on land will be an increasingly important component of polar bear conservation.

Alaska

A serological survey of Francisella tularensis exposure in wildlife on the Arctic Coastal Plain of Alaska

Tularemia is an infectious zoonotic disease caused by one of several subspecies of Francisella tularensis bacteria. Infections by F. tularensis are common throughout the northern hemisphere and have been detected in more than 250 wildlife species. In Alaska, US, where the pathogen was first identified in 1938, studies have identified F. tularensis antibodies in a diverse suite of taxa, including insects, birds, and mammals. However, few such investigations have been conducted recently and knowledge about the current distribution and disease ecology of F. tularensis is limited, particularly in Arctic Alaska, an area undergoing rapid environmental changes from climate warming. To help address these information gaps and provide insights about patterns of exposure among wildlife, we assessed the seroprevalence of F. tularensis antibodies in mammals and tundra-nesting geese from the Arctic Coastal Plain of Alaska, 2014–17. With a commercially available slide agglutination test, we detected antibodies in 14.7% of all individuals sampled ( n =722), with titers ranging from 1:20 to 1:320. We detected significant differences in seroprevalence between family groups, with Canidae (foxes, Vulpes spp.) and Sciuridae (Arctic ground squirrel, Spermophilus parryii ) having the highest seroprevalence at 21.5% and 33.3%, respectively. Mean seroprevalence for Ursidae (polar bears, Ursus maritimus ) was 13.3%, whereas Cervidae (caribou, Rangifer tarandus ) had comparatively low seroprevalence at 6.5%. Antibodies were detected in all Anatidae species sampled, with Black Brant ( Branta bernicla nigricans ) having the highest seroprevalence at 13.6%. The detection of F. tularensis antibodies across multiple taxa from the Arctic Coastal Plain and its nearshore marine region provides evidence of exposure to this pathogen throughout the region and highlights the need for renewed surveillance in Alaska.

Alaska

Using in situ/ex situ research collaborations to support polar bear conservation

A warming Arctic threatens the long-term persistence of polar bears ( Ursus maritimus ) in the wild. Historically, little collaboration existed between the in situ and ex situ polar bear scientific communities. However, for the past decade, zoo professionals, government agencies, and non-governmental organizations (NGO’s) have partnered to leverage resources and expertise with the goal of addressing science needs and enhancing polar bear conservation efforts. We discuss examples where collaborative efforts have addressed polar bear research needs and highlight opportunities for future integrated in situ and ex situ research.

Book chapter

Diet-driven mercury contamination is associated with polar bear gut microbiota

The gut microbiota may modulate the disposition and toxicity of environmental contaminants within a host but, conversely, contaminants may also impact gut bacteria. Such contaminant-gut microbial connections, which could lead to alteration of host health, remain poorly known and are rarely studied in free-ranging wildlife. The polar bear ( Ursus maritimus ) is a long-lived, wide-ranging apex predator that feeds on a variety of high trophic position seal and cetacean species and, as such, is exposed to among the highest levels of biomagnifying contaminants of all Arctic species. Here, we investigate associations between mercury (THg; a key Arctic contaminant), diet, and the diversity and composition of the gut microbiota of polar bears inhabiting the southern Beaufort Sea, while accounting for host sex, age class and body condition. Bacterial diversity was negatively associated with seal consumption and mercury, a pattern seen for both Shannon and Inverse Simpson alpha diversity indices (adjusted R 2 = 0.35, F 1,18 = 8.00, P = 0.013 and adjusted R 2 = 0.26, F 1,18 = 6.04, P = 0.027, respectively). No association was found with sex, age class or body condition of polar bears. Bacteria known to either be involved in THg methylation or considered to be highly contaminant resistant, including Lactobacillales, Bacillales and Aeromonadales, were significantly more abundant in individuals that had higher THg concentrations. Conversely, individuals with higher THg concentrations showed a significantly lower abundance of Bacteroidales, a bacterial order that typically plays an important role in supporting host immune function by stimulating intraepithelial lymphocytes within the epithelial barrier. These associations between diet-acquired mercury and microbiota illustrate a potentially overlooked outcome of mercury accumulation in polar bears.

Scientific Reports

Effects of sea ice decline and summer land use on polar bear home range size in the Beaufort Sea

Animals responding to habitat loss and fragmentation may increase their home ranges to offset declines in localized resources or they may decrease their home ranges and switch to alternative resources. In many regions of the Arctic, polar bears ( Ursus maritimus ) exhibit some of the largest home ranges of any quadrupedal mammal. Polar bears are presently experiencing a rapid decline in Arctic sea ice extent and a change in sea ice composition. For the Southern Beaufort Sea subpopulation of polar bears, this has resulted in a divergent movement pattern where most of the subpopulation remains on the sea ice in the summer melt season while the remainder move to land. We evaluated the effects of summer land use and maternal denning on the annual and seasonal utilization distribution size (i.e., home range) of adult female polar bears in the Southern Beaufort Sea subpopulation over 30 yr (1986–2016) during a period of rapid sea ice decline. For bears that remained on the summer sea ice, model-derived mean annual utilization distributions were 64% larger in 1999–2016 ( x ̄ "> 𝑥 ̄ = 176,000 km 2 ) relative to 1986–1998 ( x ̄ "> 𝑥 ̄ = 107,000 km 2 ). This increase was primarily driven by increases in summer utilization distributions that encompassed increased amounts of open water and decreased amounts of preferred sea ice. The mean centroid of summer utilization distributions for bears that remained on the sea ice was 193 km further north-northeast in 1999–2016. In contrast, bears that summered on land during 1999–2016 exhibited 88% smaller mean annual utilization distribution sizes ( x ̄ "> 𝑥 ̄ = 22,000 km 2 ) relative to bears that remained on the summer sea ice during the same period. Our findings highlight the impacts of sea ice declines on polar bear space use and the increasing importance of land as an alternative summer refuge.

Alaska