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Uncertainties in forecasting the response of polar bears to global climate change

Several sources of uncertainty affect how precisely the future status of polar bears ( Ursus maritimus ) can be forecasted. Foremost are unknowns about the future levels of global greenhouse gas emissions, which could range from an unabated increase to an aggressively mitigated reduction. Uncertainties also arise because different climate models project different amounts and rates of future warming (and sea ice loss)—even for the same emission scenario. There are also uncertainties about how global warming could affect the Arctic Ocean’s food web, so even if climate models project the presence of sea ice in the future, the availability of polar bear prey is not guaranteed. Under a worst-case emission scenario in which rates of greenhouse gas emissions continue to rise unabated to century’s end, the uncertainties about polar bear status center on a potential for extinction. If the species were to persist, it would likely be restricted to a high-latitude refugium in northern Canada and Greenland—assuming a food web also existed with enough accessible prey to fuel weight gains for surviving onshore during the most extreme years of summer ice melt. On the other hand, if emissions were to be aggressively mitigated at the levels proposed in the Paris Climate Agreement, healthy polar bear populations would probably continue to occupy all but the most southern areas of their contemporary summer range. While polar bears have survived previous warming phases—which indicate some resiliency to the loss of sea ice habitat—what is certain is that the present pace of warming is unprecedented and will increasingly expose polar bears to historically novel stressors.

Book chapter

Monitoring the welfare of polar bear populations in a rapidly changing Arctic

Most programs for monitoring the welfare of wildlife populations support efforts aimed at reaching discrete management objectives, like mitigating conflict with humans. While such programs can be effective, their limited scope may preclude systemic evaluations needed for large-scale conservation initiatives, like the recovery of at-risk species. We discuss select categories of metrics that can be used to monitor how polar bears ( Ursus maritimus ) are responding to the primary threat to their long-term persistence—loss of sea ice habitat due to the unabated rise in atmospheric greenhouse gas (GHG; e.g., CO 2 ) concentrations—that can also provide information on ecosystem function and health. Monitoring key aspects of polar bear population dynamics, spatial behavior, health and resiliency can provide valuable insight into ecosystem state and function, and could be a powerful tool for achieving Arctic conservation objectives, particularly those that have transnational policy implications.

Book chapter

Recent observations of intraspecific predation and cannibalism among polar bears in the southern Beaufort Sea

Intraspecific killing has been reported among polar bears (Ursus maritimus), brown bears (U. arctos), and black bears (U. americanus). Although cannibalism is one motivation for such killings, the ecological factors mediating such events are poorly understood. Between 24 January and 10 April 2004, we confirmed three instances of intraspecific predation and cannibalism in the Beaufort Sea. One of these, the first of this type ever reported for polar bears, was a parturient female killed at her maternal den. The predating bear was hunting in a known maternal denning area and apparently discovered the den by scent. A second predation event involved an adult female and cub recently emerged from their den, and the third involved a yearling male. During 24 years of research on polar bears in the southern Beaufort Sea region of northern Alaska and 34 years in northwestern Canada, we have not seen other incidents of polar bears stalking, killing, and eating other polar bears. We hypothesize that nutritional stresses related to the longer ice-free seasons that have occurred in the Beaufort Sea in recent years may have led to the cannibalism incidents we observed in 2004. ?? Springer-Verlag 2006.

Polar Biology

Population-level resource selection by sympatric brown and American black bears in Alaska

Distribution theory predicts that for two species living in sympatry, the subordinate species would be constrained from using the most suitable resources (e.g., habitat), resulting in its use of less suitable habitat and spatial segregation between species. We used negative binomial generalized linear mixed models with fixed effects to estimate seasonal population-level resource selection at two spatial resolutions for female brown bears ( Ursus arctos ) and female American black bears ( U. americanus ) in southcentral Alaska during May–September 2000. Black bears selected areas occupied by brown bears during spring which may be related to spatially restricted (i.e., restricted to low elevations) but dispersed or patchy availability of food. In contrast, black bears avoided areas occupied by brown bears during summer. Brown bears selected areas near salmon streams during summer, presumably to access spawning salmon. Use of areas with high berry production by black bears during summer appeared in response to avoidance of areas containing brown bears. Berries likely provided black bears a less nutritious, but adequate food source. We suggest that during summer, black bears were displaced by brown bears, which supports distribution theory in that black bears appeared to be partially constrained from areas containing salmon, resulting in their use of areas containing less nutritious forage. Spatial segregation of brown and American black bears apparently occurs when high-quality resources are spatially restricted and alternate resources are available to the subordinate species. This and previous work suggest that individual interactions between species can result in seasonal population-level responses.

Polar Biology

Plasticity in physiological condition of female brown bears across diverse ecosystems

Variation in life history strategies facilitates the near global distribution of mammals by expanding realized niche width. We investigated physiological plasticity in the spring body composition of adult female brown bears ( Ursus arctos ) across 4 diverse Alaskan ecosystems. Brown bears are a highly intelligent omnivore with a historic range spanning much of North America, Europe, and Asia. We hypothesized that body mass, fat mass, lean mass, and total caloric content would increase across populations with increasing food resource availability. Throughout their range, brown bears enter a period of torpor during winter months, decreasing their metabolic rate as an adaptation to this period of reduced food availability. They also give birth to and nourish offspring during this time. Due to this specific life history strategy, we further hypothesized that proportional body fat and the proportion of total calories derived from fat would be consistent across populations. Our results supported our first hypothesis: body, fat, and lean masses, and caloric content of bears across populations increased with the quality and abundance of available food. However, the proportional body fat content and proportion of calories from fat differed across populations indicating population-specific strategies to meet the demands of reduced seasonal food availability, offspring production and rearing, and climate as well as some plasticity to respond to environmental change or ecosystem perturbations. Investigations of body condition and energetics benefit from combined assessments of absolute, proportional, and caloric metrics to understand the nuances of brown bear physiological dynamics across and within populations.

Alaska

Fatty acid profiles of feeding and fasting bears: Estimating calibration coefficients, the timeframe of diet estimates, and selective mobilization during hibernation

Accurate information on diet composition is central to understanding and conserving carnivore populations. Quantitative fatty acid signature analysis (QFASA) has emerged as a powerful tool for estimating the diets of predators, but ambiguities remain about the timeframe of QFASA estimates and the need to account for species-specific patterns of metabolism. We conducted a series of feeding experiments with four juvenile male brown bears ( Ursus arctos ) to (1) track the timing of changes in adipose tissue composition and QFASA diet estimates in response to a change in diet and (2) quantify the relationship between consumer and diet FA composition (i.e., determine “calibration coefficients”). Bears were fed three compositionally distinct diets for 90–120 days each. Two marine-based diets were intended to approximate the lipid content and composition of the wild diet of polar bears ( U. maritimus ). Bear adipose tissue composition changed quickly in the direction of the diet and showed evidence of stabilization after 60 days. During hibernation, FA profiles were initially stable but diet estimates after 10 weeks were sensitive to calibration coefficients. Calibration coefficients derived from the marine-based diets were broadly similar to each other and to published values from marine-fed mink ( Mustela vison ), which have been used as a model for free-ranging polar bears. For growing bears on a high-fat diet, the temporal window for QFASA estimates was 30–90 days. Although our results reinforce the importance of accurate calibration, the similarities across taxa and diets suggest it may be feasible to develop a generalized QFASA approach for mammalian carnivores.

Journal of Comparative Physiology B: Biochemical,

Detecting grizzly bear use of ungulate carcasses using global positioning system telemetry and activity data

Global positioning system (GPS) wildlife collars have revolutionized wildlife research. Studies of predation by free-ranging carnivores have particularly benefited from the application of location clustering algorithms to determine when and where predation events occur. These studies have changed our understanding of large carnivore behavior, but the gains have concentrated on obligate carnivores. Facultative carnivores, such as grizzly/brown bears ( Ursus arctos ), exhibit a variety of behaviors that can lead to the formation of GPS clusters. We combined clustering techniques with field site investigations of grizzly bear GPS locations ( n = 732 site investigations; 2004–2011) to produce 174 GPS clusters where documented behavior was partitioned into five classes (large-biomass carcass, small-biomass carcass, old carcass, non-carcass activity, and resting). We used multinomial logistic regression to predict the probability of clusters belonging to each class. Two cross-validation methods—leaving out individual clusters, or leaving out individual bears—showed that correct prediction of bear visitation to large-biomass carcasses was 78–88%, whereas the false-positive rate was 18–24%. As a case study, we applied our predictive model to a GPS data set of 266 bear-years in the Greater Yellowstone Ecosystem (2002–2011) and examined trends in carcass visitation during fall hyperphagia (September–October). We identified 1997 spatial GPS clusters, of which 347 were predicted to be large-biomass carcasses. We used the clustered data to develop a carcass visitation index, which varied annually, but more than doubled during the study period. Our study demonstrates the effectiveness and utility of identifying GPS clusters associated with carcass visitation by a facultative carnivore.

Greater Yellowstone Ecosystem

Habitat degradation affects the summer activity of polar bears

Understanding behavioral responses of species to environmental change is critical to forecasting population-level effects. Although climate change is significantly impacting species’ distributions, few studies have examined associated changes in behavior. Polar bear ( Ursus maritimus ) subpopulations have varied in their near-term responses to sea ice decline. We examined behavioral responses of two adjacent subpopulations to changes in habitat availability during the annual sea ice minimum using activity data. Location and activity sensor data collected from 1989 to 2014 for 202 adult female polar bears in the Southern Beaufort Sea (SB) and Chukchi Sea (CS) subpopulations were used to compare activity in three habitat types varying in prey availability: (1) land; (2) ice over shallow, biologically productive waters; and (3) ice over deeper, less productive waters. Bears varied activity across and within habitats with the highest activity at 50–75% sea ice concentration over shallow waters. On land, SB bears exhibited variable but relatively high activity associated with the use of subsistence-harvested bowhead whale carcasses, whereas CS bears exhibited low activity consistent with minimal feeding. Both subpopulations had fewer observations in their preferred shallow-water sea ice habitats in recent years, corresponding with declines in availability of this substrate. The substantially higher use of marginal habitats by SB bears is an additional mechanism potentially explaining why this subpopulation has experienced negative effects of sea ice loss compared to the still-productive CS subpopulation. Variability in activity among, and within, habitats suggests that bears alter their behavior in response to habitat conditions, presumably in an attempt to balance prey availability with energy costs.

Oecologia

Phenotypic plasticity and climate change: Can polar bears respond to longer Arctic summers with an adaptive fast?

Plasticity in the physiological and behavioural responses of animals to prolonged food shortages may determine the persistence of species under climate warming. This is particularly applicable for species that can “adaptively fast” by conserving protein to protect organ function while catabolizing endogenous tissues. Some Ursids, including polar bears ( Ursus maritimus ), adaptively fast during winter hibernation—and it has been suggested that polar bears also employ this strategy during summer. We captured 57 adult female polar bears in the Southern Beaufort Sea (SBS) during summer 2008 and 2009 and measured blood variables that indicate feeding, regular fasting, and adaptive fasting. We also assessed tissue δ13C and δ15N to infer diet, and body condition via mass and length. We found that bears on shore maintained lipid and protein stores by scavenging on bowhead whale ( Balaena mysticetus ) carcasses from human harvest, while those that followed the retreating sea ice beyond the continental shelf were food deprived. They had low ratios of blood urea to creatinine (U:C), normally associated with adaptive fasting. However, they also exhibited low albumin and glucose (indicative of protein loss) and elevated alanine aminotransferase and ghrelin (which fall during adaptive fasting). Thus, the ~ 70% of the SBS subpopulation that spends summer on the ice experiences more of a regular, rather than adaptive, fast. This fast will lengthen as summer ice declines. The resulting protein loss prior to winter could be a mechanism driving the reported correlation between summer ice and polar bear reproduction and survival in the SBS.

Oecologia

Drivers and consequences of apex predator diet composition in the Canadian Beaufort Sea

Polar bears ( Ursus maritimus ) rely on annual sea ice as their primary habitat for hunting marine mammal prey. Given their long lifespan, wide geographic distribution, and position at the top of the Arctic marine food web, the diet composition of polar bears can provide insights into temporal and spatial ecosystem dynamics related to climate-mediated sea ice loss. Polar bears with the greatest ecological constraints on diet composition may be most vulnerable to climate-related changes in ice conditions and prey availability. We used quantitative fatty acid signature analysis (QFASA) to estimate the diets of polar bears ( n = 419) in two western Canadian Arctic subpopulations (Northern Beaufort Sea and Southern Beaufort Sea) from 1999 to 2015. Polar bear diets were dominated by ringed seal ( Pusa hispida ), with interannual, seasonal, age- and sex-specific variation. Foraging area and sea ice conditions also affected polar bear diet composition. Most variation in bear diet was explained by longitude, reflecting spatial variation in prey availability. Sea ice conditions (extent, thickness, and seasonal duration) declined throughout the study period, and date of sea ice break-up in the preceding spring was positively correlated with female body condition and consumption of beluga whale ( Delphinapterus leucas ), suggesting that bears foraged on beluga whales during entrapment events. Female body condition was positively correlated with ringed seal consumption, and negatively correlated with bearded seal consumption. This study provides insights into the complex relationships between declining sea ice habitat and the diet composition and foraging success of a wide-ranging apex predator.

Northwest Territories, Yukon

Summer/fall diet and macronutrient assimilation in an Arctic predator

Free-ranging predator diet estimation is commonly achieved by applying molecular-based tracers because direct observation is not logistically feasible or robust. However, tracers typically do not represent all dietary macronutrients, which likely obscures resource use as prey proximate composition varies and tissue consumption can be specific. For example, polar bears ( Ursus maritimus ) preferentially consume blubber, yet diets have been estimated using fatty acids based on prey blubber or stable isotopes of lipid-extracted prey muscle, neither of which represent both protein and lipid macronutrient contributions. Further, additional bias can be introduced because dietary fat is known to be flexibly routed beyond short-term energy production and storage. We address this problem by simultaneously accounting for protein and lipid assimilation using carbon and nitrogen isotope compositions of lipid-containing prey muscle and blubber to infer summer/fall diet composition and macronutrient proportions from Chukchi Sea polar bear guard hair ( n = 229) sampled each spring between 2008 and 2017. Inclusion of blubber (85–95% lipid by dry mass) expanded the isotope mixing space and improved separation among prey species. Ice-associated seals, including nutritionally dependent pups, were the primary prey in summer/fall diets with lower contributions by Pacific walruses ( Odobenus rosmarus ) and whales. Percent blubber estimates confirmed preferential selection of this tissue and represented the highest documented lipid assimilation for any animal species. Our results offer an improved understanding of summer/fall prey macronutrient usage by Chukchi Sea polar bears which likely coincides with a nutritional bottleneck as the sea ice minimum is approached.

Alaska

The effects of carnivory and herbivory on the energy balance of Arctic grizzly bears

Omnivores often face tradeoffs between selecting for spatially dispersed energy-dense vertebrate prey versus densely distributed herbivorous resources that have limited energetic value per unit intake. Arctic grizzly bears ( Ursus arctos ) are large omnivores within a resource-limited ecosystem that are known to exhibit smaller body masses and occur at lower densities than grizzly bears in other regions of North America. We evaluated the energy balance of Arctic grizzly bears during a portion of the fall hyperphagic period in two ecologically differing regions on Alaska’s northern Arctic coast by monitoring mass change, food intake, activity, and energy expenditure of 12 individuals over 17–22 days. Bears in coastal areas were more carnivorous than bears in the foothills that were predominantly herbivorous and frugivorous. Carnivory was associated with greater movement, body fat, and energy expenditure and two of four carnivorous bears lost mass. Overall, the mean body fat of the bears in this study was 34% lower than other grizzly bear populations in North America in the fall. Furthermore, the bears in this study exhibited relatively small changes in body mass (x̄ = 3%, range =−2 to 11%) that were 60% lower than other grizzly bear populations which typically gain substantial mass in the fall in preparation for denning. Our results, while representing a snapshot from a small number of bears during the fall hyperphagic period, are consistent with previous studies and indicate limited availability of energy-dense food resources during this time for grizzly bears in this region of the Arctic.

Alaska

Hematology of southern Beaufort Sea polar bears (2005-2007): Biomarker for an arctic ecosystem health sentinel

Declines in sea-ice habitats have resulted in declining stature, productivity, and survival of polar bears in some regions. With continuing sea-ice declines, negative population effects are projected to expand throughout the polar bear's range. Precise causes of diminished polar bear life history performance are unknown, however, climate and sea-ice condition change are expected to adversely impact polar bear (Ursus maritimus) health and population dynamics. As apex predators in the Arctic, polar bears integrate the status of lower trophic levels and are therefore sentinels of ecosystem health. Arctic residents feed at the apex of the ecosystem, thus polar bears can serve as indicators of human health in the Arctic. Despite their value as indicators of ecosystem welfare, population-level health data for U.S. polar bears are lacking. We present hematological reference ranges for southern Beaufort Sea polar bears. Hematological parameters in southern Beaufort Sea polar bears varied by age, geographic location, and reproductive status. Total leukocytes, lymphocytes, monocytes, eosinophils, and serum immunoglobulin G were significantly greater in males than females. These measures were greater in nonlactating females ages ???5, than lactating adult females ages ???5, suggesting that females encumbered by young may be less resilient to new immune system challenges that may accompany ongoing climate change. Hematological values established here provide a necessary baseline for anticipated changes in health as arctic temperatures warm and sea-ice declines accelerate. Data suggest that females with dependent young may be most vulnerable to these changes and should therefore be a targeted cohort for monitoring in this sentinel. ?? 2010 International Association for Ecology and Health.

EcoHealth

Using gene transcription to assess ecological and anthropological stressors in brown bears

Increasingly, population- and ecosystem-level health assessments are performed using sophisticated molecular tools. Advances in molecular technology enable the identification of synergistic effects of multiple stressors on the individual physiology of different species. Brown bears ( Ursus arctos ) are an apex predator; thus, they are ideal candidates for detecting potentially ecosystem-level systemic perturbations using molecular-based tools. We used gene transcription to analyze 130 brown bear samples from three National Parks and Preserves in Alaska. Although the populations we studied are apparently stable in abundance and exist within protected and intact environments, differences in transcript profiles were noted. The most prevalent differences were among locations. The transcript patterns among groups reflect the influence of environmental factors, such as nutritional status, disease, and xenobiotic exposure. However, these profiles also likely represent baselines for each unique environment by which future measures can be made to identify early indication of population-level changes due to, for example, increasing Arctic temperatures. Some of those environmental changes are predicted to be potentially positive for brown bears, but other effects such as the manifestation of disease or indirect effects of oceanic acidification may produce negative impacts.

Alaska

Genetic diversity, effective population size, and structure among black bear populations in the Lower Mississippi Alluvial Valley, USA

Multiple small populations of American black bears Ursus americanus , including the recently delisted Louisiana black bear subspecies U. a. luteolus , occupy a fragmented landscape in the Lower Mississippi Alluvial Valley, USA (LMAV). Populations include bears native to the LMAV, bears translocated from Minnesota during the 1960s, and recently reintroduced and colonizing populations sourced from within the LMAV. We estimated population structure, gene flow, and genetic parameters important to conservation of small populations using genotypes at 23 microsatellite markers for 265 bears from seven populations. We inferred five genetic clusters corresponding to the following populations: White River and western Mississippi, Tensas River and Three Rivers, Upper Atchafalaya, Lower Atchafalaya, and Minnesota. Upper Atchafalaya was suggested as the product of Minnesota-sourced translocations, but those populations have since diverged, likely because of a founder effect followed by genetic drift and isolation. An admixture zone recently developed in northeastern Louisiana and western Mississippi between migrants from White River and Tensas River, resulting in a Wahlund effect. However, gene flow among most populations has been limited and considerable genetic differentiation accumulated (global F ST = 0.22), particularly among the three Louisiana black bear populations that existed when federal listing occurred. Consistent with previous bottlenecks, founder effects, and persisting isolation, all LMAV bear populations had low genetic diversity ( A R = 2.08–4.81; H E = 0.36–0.63) or small effective population size ( N E = 3–49). Translocating bears among populations as part of a regional genetic restoration program may help improve genetic diversity and increase effective population sizes.

Lower Mississippi Alluvial Valley

Identification of functional corridors with movement characteristics of brown bears on the Kenai Peninsula, Alaska

We identified primary habitat and functional corridors across a landscape using Global Positioning System (GPS) collar locations of brown bears (Ursus arctos). After deriving density, speed, and angular deviation of movement, we classified landscape function for a group of animals with a cluster analysis. We described areas with high amounts of sinuous movement as primary habitat patches and areas with high amounts of very directional, fast movement as highly functional bear corridors. The time between bear locations and scale of analysis influenced the number and size of corridors identified. Bear locations should be collected at intervals ???6 h to correctly identify travel corridors. Our corridor identification technique will help managers move beyond the theoretical discussion of corridors and linkage zones to active management of landscape features that will preserve connectivity. ?? 2007 Springer Science+Business Media, Inc.

Landscape Ecology

Hidden Markov model for dependent mark loss and survival estimation

Mark-recapture estimators assume no loss of marks to provide unbiased estimates of population parameters. We describe a hidden Markov model (HMM) framework that integrates a mark loss model with a Cormack–Jolly–Seber model for survival estimation. Mark loss can be estimated with single-marked animals as long as a sub-sample of animals has a permanent mark. Double-marking provides an estimate of mark loss assuming independence but dependence can be modeled with a permanently marked sub-sample. We use a log-linear approach to include covariates for mark loss and dependence which is more flexible than existing published methods for integrated models. The HMM approach is demonstrated with a dataset of black bears ( Ursus americanus ) with two ear tags and a subset of which were permanently marked with tattoos. The data were analyzed with and without the tattoo. Dropping the tattoos resulted in estimates of survival that were reduced by 0.005–0.035 due to tag loss dependence that could not be modeled. We also analyzed the data with and without the tattoo using a single tag. By not using.

Journal of Agricultural, Biological, and Environme

Movements of Yellowstone grizzly bears

Ninety-seven grizzly bears Ursus arctos horribilis were radio-located 6299 times during 1975–1987. Annual range sizes differed by sex, age, reproductive status and amount of precipitation. Females exhibited greater fidelity to seasonal and annual ranges than males. Weaned female offspring generally remained in the vicinity of the maternal range, while weaned males often made substantial movements to unexplored country. Average total home range size was 884 km 2 for females and 3757 km 2 for males. Males consistently exhibited greater indices of movement and range sizes than females. All cohorts had larger mean ranges during this study than during the period 1959–1969 when the population had access to open garbage dumps. Movements and elevational distribution of all cohorts were related to availability of whitebark pine Pinus albicaulis seeds. We hypothesized that females with cubs-of-the-year and yearlings were displaced from most productive habitats during seasons and years of limited food availability.

Idaho;Montana;Wyoming