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.     

Fasting physiology of polar bears in relation to environmental change and breeding behavior in the Beaufort Sea

We examined the use of the ratio of serum urea to serum creatinine as a physiological biomarker of fasting to monitor temporal patterns in the feeding ecology of polar bears (Ursus maritimus). Blood was collected from 436 polar bears in the eastern Beaufort Sea during April and May of 1985–1986 and 2005–2006. The proportions of polar bears fasting were 9.6% in 1985, 10.5% in 1986, 21.4% in 2005, and 29.3% in 2006. We used stepwise logistic regression analysis to evaluate factors that could influence the binary response variable of fasting or not fasting. Significant predictor variables of fasting were: the 2005 and 2006 capture years, solitary adult male bears, and adult male bears that were accompanying an estrous female. The increased number of polar bears in a physiological fasting state from all sex, age, and reproductive classes in 2005 and 2006 corresponded with broad scale changes in Arctic sea ice composition, which may have affected prey availability. The higher proportion of adult males fasting from all years was attributed to spring breeding behavior.     

Population ecology of polar bears in Davis Strait,Canada and Greenland

Until recently, the sea ice habitat of polar bears was understood to be variable, but environmental variability was considered to be cyclic or random, rather than progressive. Harvested populations were believed to be at levels where density effects were considered not significant. However, because we now understand that polar bear demography can also be influenced by progressive change in the environment, and some populations have increased to greater densities than historically lower numbers, a broader suite of factors should be considered in demographic studies and management. We analyzed 35 years of capture and harvest data from the polar bear (Ursus maritimus) subpopulation in Davis Strait, including data from a new study (2005–2007), to quantify its current demography. We estimated the population size in 2007 to be 2,158 ± 180 (SE), a likely increase from the 1970s. We detected variation in survival, reproductive rates, and age-structure of polar bears from geographic sub-regions. Survival and reproduction of bears in southern Davis Strait was greater than in the north and tied to a concurrent dramatic increase in breeding harp seals (Pagophilus groenlandicus) in Labrador. The most supported survival models contained geographic and temporal variables. Harp seal abundance was significantly related to polar bear survival. Our estimates of declining harvest recovery rate, and increasing total survival, suggest that the rate of harvest declined over time. Low recruitment rates, average adult survival rates, and high population density, in an environment of high prey density, but deteriorating and variable ice conditions, currently characterize the Davis Strait polar bears. Low reproductive rates may reflect negative effects of greater densities or worsening ice conditions. © 2013 The Wildlife Society.     

Demography and population viability of polar bears in the Gulf of Boothia, Nunavut

We estimated demographic parameters and harvest risks for polar bears ( Ursus maritimus ) inhabiting the Gulf of Boothia, Nunavut, from 1976 to 2000. We computed survival and abundance from capture–recapture and recovery data (630 marks) using a Burnham joint live–dead model implemented in program MARK. Annual mean total survival (including harvest) was 0.889 ± 0.179 (± 1 SE) for cubs, 0.883 ± 0.087 for subadults (ages 1–4), 0.919 ± 0.044 for adult females, and 0.917 ± 0.041 for adult males. Abundance in the last 3 yr of study was 1,592 ± 361 bears. Mean size of newborn litters was 1.648 ± 0.098 cubs. By age 7, 0.97 ± 0.30 of available females were producing litters. Harvest averaged 38.4 ± 4.2 bears/year in the last 5 yr of study; however, the 2002–2007 kill averaged 56.4 bears/yr. We used a harvested Population Viability Analysis (PVA) to examine impacts of increasing rates of harvest. We estimated the current population growth rate, λ _H , to be 1.025 ± 0.032. Although this suggests the population is growing, progressive environmental changes may require more frequent population inventory studies to maintain the same levels of harvest risk.     

Stable isotopes to detect food-conditioned bears and to evaluate human-bear management

We used genetic and stable isotope analysis of hair from free-ranging black bears (Ursus americanus) in Yosemite National Park, California, USA to: 1) identify bears that consume human food, 2) estimate the diets of these bears, and 3) evaluate the Yosemite human–bear management program. Specifically, we analyzed the isotopic composition of hair from bears known a priori to be food-conditioned or non-food-conditioned and used these data to predict whether bears with an unknown management status were food-conditioned (FC) or non-food-conditioned (NFC). We used a stable isotope mixing model to estimate the proportional contribution of natural foods (plants and animals) versus human food in the diets of FC bears. We then used results from both analyses to evaluate proactive (population-level) and reactive (individual-level) human–bear management, and discussed new metrics to evaluate the overall human–bear management program in Yosemite. Our results indicated that 19 out of 145 (13%) unknown bears sampled from 2005 to 2007 were food-conditioned. The proportion of human food in the diets of known FC bears likely declined from 2001–2003 to 2005–2007, suggesting proactive management was successful in reducing the amount of human food available to bears. In contrast, reactive management was not successful in changing the management status of known FC bears to NFC bears, or in reducing the contribution of human food to the diets of FC bears. Nine known FC bears were recaptured on 14 occasions from 2001 to 2007; all bears were classified as FC during subsequent recaptures, and human–bear management did not reduce the amount of human food in the diets of FC bears. Based on our results, we suggest Yosemite continue implementing proactive human–bear management, reevaluate reactive management, and consider removing problem bears (those involved in repeated bear incidents) from the population. © 2012 The Wildlife Society.     

Survival and abundance of polar bears in Alaska’s Beaufort Sea, 2001–2016

The Arctic Ocean is undergoing rapid transformation toward a seasonally ice‐free ecosystem. As ice‐adapted apex predators, polar bears (Ursus maritimus) are challenged to cope with ongoing habitat degradation and changes in their prey base driven by food‐web response to climate warming. Knowledge of polar bear response to environmental change is necessary to understand ecosystem dynamics and inform conservation decisions. In the southern Beaufort Sea (SBS) of Alaska and western Canada, sea ice extent has declined since satellite observations began in 1979 and available evidence suggests that the carrying capacity of the SBS for polar bears has trended lower for nearly two decades. In this study, we investigated the population dynamics of polar bears in Alaska''s SBS from 2001 to 2016 using a multistate Cormack–Jolly–Seber mark–recapture model. States were defined as geographic regions, and we used location data from mark–recapture observations and satellite‐telemetered bears to model transitions between states and thereby explain heterogeneity in recapture probabilities. Our results corroborate prior findings that the SBS subpopulation experienced low survival from 2003 to 2006. Survival improved modestly from 2006 to 2008 and afterward rebounded to comparatively high levels for the remainder of the study, except in 2012. Abundance moved in concert with survival throughout the study period, declining substantially from 2003 and 2006 and afterward fluctuating with lower variation around an average of 565 bears (95% Bayesian credible interval [340, 920]) through 2015. Even though abundance was comparatively stable and without sustained trend from 2006 to 2015, polar bears in the Alaska SBS were less abundant over that period than at any time since passage of the U.S. Marine Mammal Protection Act. The potential for recovery is likely limited by the degree of habitat degradation the subpopulation has experienced, and future reductions in carrying capacity are expected given current projections for continued climate warming.     

Investigating the potential use of aerial line transect surveys for estimating polar bear abundance in sea ice habitats: A case study for the Chukchi Sea

The expense of traditional capture‐recapture methods, interest in less invasive survey methods, and the circumpolar decline of polar bear (Ursus maritimus) habitat require evaluation of alternative methods for monitoring polar bear populations. Aerial line transect distance sampling (DS) surveys are thought to be a promising monitoring tool. However, low densities and few observations during a survey can result in low precision, and logistical constraints such as heavy ice and fuel and safety limitations may restrict survey coverage. We used simulations to investigate the accuracy and precision of, DS for estimating polar bear abundance in sea ice habitats, using the Chukchi Sea subpopulation as an example. Simulation parameters were informed from a recent pilot survey. Predictions from a resource selection model were used for stratification, and we compared two ratio estimators to account for areas that cannot be sampled. The ratio estimator using predictions of resource selection by polar bears allowed for extrapolation beyond sampled areas and provided results with low bias and CVs ranging from 21% to 36% when abundance was >1,000. These techniques could be applied to other DS surveys to allocate effort and potentially extrapolate estimates to include portions of the landscape that are logistically impossible to survey.     

Increased Arctic sea ice drift alters adult female polar bear movements and energetics

Recent reductions in thickness and extent have increased drift rates of Arctic sea ice. Increased ice drift could significantly affect the movements and the energy balance of polar bears (Ursus maritimus) which forage, nearly exclusively, on this substrate. We used radio‐tracking and ice drift data to quantify the influence of increased drift on bear movements, and we modeled the consequences for energy demands of adult females in the Beaufort and Chukchi seas during two periods with different sea ice characteristics. Westward and northward drift of the sea ice used by polar bears in both regions increased between 1987–1998 and 1999–2013. To remain within their home ranges, polar bears responded to the higher westward ice drift with greater eastward movements, while their movements north in the spring and south in fall were frequently aided by ice motion. To compensate for more rapid westward ice drift in recent years, polar bears covered greater daily distances either by increasing their time spent active (7.6%–9.6%) or by increasing their travel speed (8.5%–8.9%). This increased their calculated annual energy expenditure by 1.8%–3.6% (depending on region and reproductive status), a cost that could be met by capturing an additional 1–3 seals/year. Polar bears selected similar habitats in both periods, indicating that faster drift did not alter habitat preferences. Compounding reduced foraging opportunities that result from habitat loss; changes in ice drift, and associated activity increases, likely exacerbate the physiological stress experienced by polar bears in a warming Arctic.     

Determination of vitamins D,A, and E in sera and vitamin D in milk from captive and free-ranging polar bears (Ursus maritimus), and 7-dehydrocholesterol levels in skin from captive polar bears

The difference between serum levels from 36 captive and 56 free-ranging polar bears (Ursus maritimus) for 25-hydroxyvitamin D (25-OH-D) was found not to be significant (mean ± SD = 348 ± 215 nmol/L [captive], 360 ± 135 nmol/L [free-ranging], t = 0.30, df = 52.8, P = 0.76), whereas the difference for retinol and α-tocopherol was significant (retinol, 1.37 ± 0.67 μmol/L [captive] 1.89 ± 0.63 μmol/L [free-ranging], t = 3.88, df = 72.4, P <0.001, α-tocopherol, 18.56 ± 18.56 μmol/L [captive], 48.76 ± 13.92 μmol/L [free-ranging], t = 7.85, df = 61.9, P < 0.001). Due to the high fat content in the polar bear diet, seal blubber may be the source of these fat-soluble vitamins. Six skin biopsies were analyzed from captive polar bears at the Denver Zoological Gardens for 7-dehydrocholesterol levels and found to contain 0.11 ± 0.03 nmol/cm². This finding also helps to support the contention that the source of vitamin D for polar bears may be ingestion and not cutaneous production. Vitamin D content in the milk from one captive sow in the den (0.14 nmol/g) and 10 free-ranging sows with cubs of the year out on the ice pack (0.0042 ± 0.0073 nmol/g) were also evaluated. It would be helpful to evaluate additional milk samples from denning and non-denning sows with cubs to see whether vitamin D content varies according to the stage of lactation. Zoo Biol 17:285–293, 1998. © 1998 Wiley- Liss, Inc.     

Heritability of body size in the polar bears of Western Hudson Bay

Among polar bears (Ursus maritimus), fitness is dependent on body size through males’ abilities to win mates, females’ abilities to provide for their young and all bears’ abilities to survive increasingly longer fasting periods caused by climate change. In the Western Hudson Bay subpopulation (near Churchill, Manitoba, Canada), polar bears have declined in body size and condition, but nothing is known about the genetic underpinnings of body size variation, which may be subject to natural selection. Here, we combine a 4449‐individual pedigree and an array of 5,433 single nucleotide polymorphisms (SNPs) to provide the first quantitative genetic study of polar bears. We used animal models to estimate heritability (h²) among polar bears handled between 1966 and 2011, obtaining h² estimates of 0.34–0.48 for strictly skeletal traits and 0.18 for axillary girth (which is also dependent on fatness). We genotyped 859 individuals with the SNP array to test for marker–trait association and combined p‐values over genetic pathways using gene‐set analysis. Variation in all traits appeared to be polygenic, but we detected one region of moderately large effect size in body length near a putative noncoding RNA in an unannotated region of the genome. Gene‐set analysis suggested that variation in body length was associated with genes in the regulatory cascade of cyclin expression, which has previously been associated with body size in mice. A greater understanding of the genetic architecture of body size variation will be valuable in understanding the potential for adaptation in polar bear populations challenged by climate change.     

Effects of sea ice extent and food availability on spatial and temporal distribution of polar bears during the fall open-water period in the Southern Beaufort Sea

We investigated the relationship between sea ice conditions, food availability, and the fall distribution of polar bears (Ursus maritimus) in terrestrial habitats of the Southern Beaufort Sea via weekly aerial surveys in 2000–2005. Aerial surveys were conducted weekly during September and October along the Southern Beaufort Sea coastline and barrier islands between Barrow and the Canadian border to determine polar bear density on land. The number of bears on land both within and among years increased when sea-ice was retreated furthest from the shore. However, spatial distribution also appeared to be related to the availability of subsistence-harvested bowhead whale (Balaena mysticetus) carcasses and the density of ringed seals (Phoca hispida) in offshore waters. Our results suggest that long-term reductions in sea-ice could result in an increasing proportion of the Southern Beaufort Sea polar bear population coming on land during the fall open-water period and an increase in the amount of time individual bears spend on land.     

Unusual abundance of macrobenthos and biological invasions in the Chukchi Sea

The data from the expedition of the program RUSALCA conducted in 2004 showed unexpectedly high quantitative indices of macrobenthos in the southeastern Chukchi Sea. Extensive areas of the bottom northwest of the Bering Strait were dominated by the bivalve Macoma calcarea. The greatest biomass of benthos in Macoma-dominated areas was 4232 g/m² with an average of 1382 g/m² for the investigated region. Such a high biomass of soft-bottom communities, which is extremely uncommon even in the temperature regions of the oceans, is reported for the Arctic for the first time. The long-term existence (more than 70 years) of highly productive benthic communities dominated by Macoma calcarea in one and the same area of the Chukchi Sea can most likely be attributed to gyres, which constantly arise in the region northwest of the Bering Strait. These cyclonic gyres carry nutrient-rich bottom water to the surface and hinder larval transport away from mother populations. They also keep and concentrate major food sources of benthos (live and dead phyto-and zooplankton and fecal pellets) over the benthic community locations. Most likely, a significant proportion of the primary production in the southeastern Chukchi Sea is used by benthos within the investigated Macoma community. Findings of three relatively large warm-water Pacific species near Point Hope in the Chukchi Sea are probably indicative of the progressive climate warming during the last century.     

Ringed seal (Pusa hispida) tooth annuli as an index of reproduction in the Beaufort Sea

Multi-decadal time-series of biological indices that reflect the state of a population are rare in ecological studies, but invaluable for assessing environmental regulation of population dynamics. We utilized canine teeth extracted from ringed seals (Pusa hispida) killed by polar bears (Ursus maritimus) in the Beaufort Sea, Canada, in 1985–2011, to obtain widths of annual growth layers in the cementum. Canine teeth for 75 individuals were measured and compared across years using a proportional width index (PWI) spanning 1965–2007. PWI was positively correlated with ringed seal ovulation rate obtained independently from other studies and was significantly lower than normal during ringed seal reproductive declines in 1974–75, 1984–87, 1991–93, and 2004–05, suggesting that PWI reflects ringed seal reproductive capacity. The PWI was also examined against climatic and sea ice factors to assess environmental regulation of ringed seal reproduction. Results suggest that ringed seals benefit from cyclonic circulation regimes in the Beaufort Sea, and an earlier breakup of sea ice in summer that may positively influence the quality and quantity of food during the open water season. Results highlight how cementum annuli in the canine teeth of ringed seals can provide an index of body state and linkages to sea ice conditions. Canine teeth from ringed seals can function as a means to monitor the effects of past Arctic marine variability on area-specific populations for which there are few independent empirical data.     

Dietary habits of polar bears in Foxe Basin,Canada: possible evidence of a trophic regime shift mediated by a new top predator

Polar bear (Ursus maritimus) subpopulations in several areas with seasonal sea ice regimes have shown declines in body condition, reproductive rates, or abundance as a result of declining sea ice habitat. In the Foxe Basin region of Nunavut, Canada, the size of the polar bear subpopulation has remained largely stable over the past 20 years, despite concurrent declines in sea ice habitat. We used fatty acid analysis to examine polar bear feeding habits in Foxe Basin and thus potentially identify ecological factors contributing to population stability. Adipose tissue samples were collected from 103 polar bears harvested during 2010–2012. Polar bear diet composition varied spatially within the region with ringed seal (Pusa hispida) comprising the primary prey in northern and southern Foxe Basin, whereas polar bears in Hudson Strait consumed equal proportions of ringed seal and harp seal (Pagophilus groenlandicus). Walrus (Odobenus rosmarus) consumption was highest in northern Foxe Basin, a trend driven by the ability of adult male bears to capture large‐bodied prey. Importantly, bowhead whale (Balaena mysticetus) contributed to polar bear diets in all areas and all age and sex classes. Bowhead carcasses resulting from killer whale (Orcinus orca) predation and subsistence harvest potentially provide an important supplementary food source for polar bears during the ice‐free period. Our results suggest that the increasing abundance of killer whales and bowhead whales in the region could be indirectly contributing to improved polar bear foraging success despite declining sea ice habitat. However, this indirect interaction between top predators may be temporary if continued sea ice declines eventually severely limit on‐ice feeding opportunities for polar bears.     

Abundance and distribution of toxic Alexandrium tamarense resting cysts in the sediments of the Chukchi Sea and the eastern Bering Sea

Abundance and distribution of the toxic dinoflagellate Alexandrium tamarense species complex resting cyst were investigated in the eastern Bering Sea and the Chukchi Sea for the first time. Sediment samples (top 0–3 cm depth) were collected from the continental shelf of the eastern Bering Sea (17 stations) and the Chukchi Sea (13 stations) together with a long core sample (top 0–21 cm depth) from one station in the Chukchi Sea during 2009–2012. The cysts were enumerated using the primuline staining method. Species identification of the cysts was carried out with multiplex PCR assay and the plate morphology of vegetative cells germinated from cysts in the both areas. Alexandrium cysts were widely detected in the both areas, ranging from not detected (<1 cysts cm⁻³) to 835 cysts cm⁻³ wet sediment in the eastern Bering Sea and from not detected (<1 cysts cm⁻³) to 10,600 cysts cm⁻³ in the Chukchi Sea, and all isolated cysts were genetically and morphologically identified as the North American clade A. tamarense. Their cysts were mainly distributed in the shallow continental shelf where the water depth was less than 100 m in both areas. The cysts were detected from the deep layer (18–21 cm depth of sediment core) of the long core sample. The present study confirmed the abundant existence of A. tamarense with wide range of distribution in these areas. This fact suggests that A. tamarense vegetative cells have appeared in the water column in the both areas. Furthermore, these abundant cyst depositions indicate that this species originally distributed in the Arctic and subarctic regions and well adapted to the environments in the marginal ice zone.     

25‐OH‐D3, calcium,phosphorus, and alkaline phosphatase content in sera from captive polar bear (Ursus maritimus) cubs at the Denver Zoological Foundation

After the development of rickets in two hand‐reared polar bear (Ursus maritimus) cubs at the Denver Zoological Foundation we initiated a project opportunistically evaluating sera 25‐OH‐D₃, calcium, phosphorus, and alkaline phosphatase levels in seven mother‐reared polar bear cubs. From one to three serum samples were obtained while the cub was still in the den with no access to natural sunlight and one after the cub exited the den. The sera 25‐OH‐D₃ increased as the cubs grew (108±37 nmol/L [n=7] while in den, 184.6±48.3 nmol/L [n=7] out of the den) but were substantially lower then captive adults (348±215 nmol/L [n=36]) or free‐ranging adults (360±135 nmol/L [n=56]). We found that age was correlated significantly with 25‐OH‐D₃ levels but den status (in or out) was not significant. Sera calcium and phosphorus values were highest in denned cubs (calcium, 2.9±0.4 mmol/L [n=13]; phosphorus, 3.1±0.5 mmol/L [n=13]) and then declined as the cubs matured and left the den (calcium, 2.5±0.1 mmol/L [n=7]; phosphorus, 2.4±0.4 mmol/L [n=7]). The enzyme alkaline phosphatase (presumably the bone isoenzyme) was high in denned growing cubs (831±426 IU/L [n=13]) and then declined gradually as the cubs aged and exited the den (96±83 IU/L [n=7]). Zoo Biol 0:1–7, 2007. © 2007 Wiley‐Liss, Inc.