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Polar bears (Ursus maritimus) in the southern Beaufort Sea experience long annual periods when preferred seal prey are scarce or are unavailable. Consumption of bowhead whale (Balaena mysticetus) carcasses from native Alaskan subsistence hunting is increasingly common for onshore polar bears, yet the energetic consequences of this consumption remain unclear. We use data on bears captured repeatedly over periods that encompassed autumn and winter, combined with calculations, to show that adult female bears likely consume an average of at least 4 seal equivalents during both autumn and winter periods and that considerable variation in energy intake exists across individual bears. We further show that subsistence-caught whale carcasses provide an upper threshold of > 4000 seal equivalents, which could potentially meet mean consumption needs of ~ 80% of the southern Beaufort Sea bear subpopulation during autumn and winter periods. Finally, we modify an existing model to show that observed mass changes over autumn and winter could substantially alter spring foraging habitat choice by females with cubs and the chance that a female with reduced energy reserves would abort a pregnancy or abandon cubs in favor of increasing her own survival; these behaviors could potentially influence population vital rates. Our study highlights the importance of mass dynamics over the autumn and winter months, points to the need for additional data on foraging and energetics over this period, and indicates that the recent declines in polar bear body condition in some subpopulations could have complex effects on reproduction.

Changes in the nutritional status of free-ranging animals have a strong influence on individual fitness, yet it remains challenging to monitor longitudinally. Nitrogen (δ15N) and carbon (δ13C) isotope values measured chronologically along the length of metabolically inert keratinous tissues can be used as a nutritional biomarker to retrospectively reconstruct the foraging ecology and eco-physiology of consumers. We quantitatively describe the physiological effects of fasting on amino acid metabolism using sequentially measured bulk tissue and amino acid δ15N values along the length of whiskers sampled from free-ranging juvenile, subadults, adult female, and male southern elephant seals (SES; Mirounga leonina) on Marion Island in the Southern Ocean. For both juveniles and adult females, whisker segments representing fasting had significantly higher bulk tissue δ15N values of 0.6 ± 0.5‰ and 1.3–1.8‰, respectively, in comparison to segments unaffected by fasting. We also found a large increase (2–6‰) in δ15N values for most glucogenic amino acids and a simultaneous depletion (2–3‰) of alanine in segments reflecting fasting, which enabled us to accurately predict (74%) the nutritional status of our model species. We hypothesize that the glucose-alanine cycle is the mechanism driving the observed depletion of alanine δ15N values during fasting. We demonstrated that keratinaceous tissues can be used as a longitudinal nutritional biomarker to detect changes in the nitrogen balance of an individual. Moreover, it is evident that physiological factors have an important influence on tissue δ15N values and can lead to erroneous bulk tissue or amino acid isotope-based reconstructions of foraging habits.

Many wildlife species are live captured, sampled, and released; for polar bears (

Nitrogen isotope (δ

The dynamics of animal body water and metabolism are integral aspects of biological function but are difficult to measure, particularly in free-ranging individuals. We demonstrate a new method to estimate inputs to body water via analysis of Δ17O, a measure of 17O/16O relative to 18O/16O. Animal body water is primarily a mixture of drinking or food water (meteoric water; Δ17O ≈ 0.030 per mille [‰]) and metabolic water synthesized from atmospheric oxygen (Δ17O ≈ –0.450‰). Greater drinking or food water intake should increase Δ17O toward 0.030‰, whereas greater metabolic rate should decrease Δ17O toward –0.450‰. We found that wild mammal Δ17O values generally increased with body mass, consistent with both a decline in mass-specific metabolic rate and an increase in water intake. Captive mouse (Peromyscus maniculatus) Δ17O values were higher than predicted but exhibited the expected relative change based on metabolic rate and water intake. Measurements of Δ17O may enable novel ecophysiological studies.

Stable isotopes are important ecological tools, because the carbon and nitrogen isotopic composition of consumer tissue reflects the diet. Measurements of isotopes of individual amino acids can disentangle the effects of consumer physiology from spatiotemporal variation in dietary isotopic values. However, this approach requires knowledge of assimilation patterns of dietary amino acids. We reared leopard sharks (Triakis semifasciata) on diets of squid (Loligo opalescens; 1250 days; control sharks) or squid then tilapia (Oreochromis sp.; switched at 565 days; experimental sharks) to evaluate consumer-diet discrimination factors for amino acids in muscle tissue. We found that control sharks exhibited lower nitrogen isotope discrimination factors (∆15N) than most previous consumer studies, potentially because of urea recycling. Control sharks also had large carbon isotope discrimination factors (∆13C) for three essential amino acids, suggesting microbial contributions or fractionation upon assimilation. Compared to controls, experimental sharks exhibited higher ∆13C values for four amino acids and ∆15N values for seven amino acids, corresponding with differences between diets in δ13C and δ15N values. This suggests that not all amino acids in experimental sharks had reached steady state, contrary to the conclusion of a bulk isotope study of these sharks. Our results imply that (1) the magnitude of a shift in dietary δ13C and δ15N values temporarily influences the appearance of discrimination factors; (2) slow turnover of amino acid isotopes in elasmobranch muscle precludes inferences about seasonal dietary changes; (3) elasmobranch discrimination factors for amino acids may be affected by urea recycling and microbial contributions of amino acids.

The sloth bear (Melursus ursinus) is one of the least studied bears. Important aspects of sloth bear biology and ecology, such as reproductive physiology and behavior, are largely unknown. Increased scent-marking by anogenital rubbing during breeding season has been recorded in other bear species. We studied the genital rubbing behavior of 37 captive female sloth bears (2–18 yr of age) at the Agra Bear Rescue Facility, India, for 4 breeding seasons over a period of 3.5 years (1 Jun 2015 to 31 Dec 2018). Data on changes in vulva visibility and presence of genital rubbing behavior were collected daily during the breeding period and twice per week for rest of the year, throughout the study period. Vulva visibility was scored as 0 (not visible), 1 (slightly visible), and 2 (fully visible), and a female was considered to be in estrus if the vulva was slightly or fully visible. Presence of genital rubbing was recorded as 1 and its absence as 0. Occurrence of genital rubbing coincided with estrus, as defined by vulva visibility scores. Statistical analysis indicated that female age and the number of males with physical proximity (i.e., in the same enclosure) were significantly correlated with the occurrence of genital rubbing behavior. The number of females in physical proximity and the number of females in the vicinity without physical proximity (i.e., not in the same enclosure but sharing a fence) did not significantly affect this behavior. The results of our study suggest that the genital rubbing behavior by female sloth bears in estrus is likely a form of scent-marking, serving a communicative function, and could be influenced by male presence. This behavior may be a key factor in attracting a mate during the breeding season in the wild.

Rationale Ecologists increasingly determine the δ15 N values of amino acids (AA) in animal tissue; "source" AA typically exhibit minor variation between diet and consumer, while "trophic" AA have increased δ15 N values in consumers. Thus, trophic-source δ15 N offsets (i.e., Δ15 NT-S ) reflect trophic position in a food web. However, even minor variations in δ15 Nsource AA values may influence the magnitude of offset that represents a trophic step, known as the trophic discrimination factor (i.e., TDFT-S ). Diet digestibility and protein content can influence the δ15 N values of bulk animal tissue, but the effects of these factors on AA Δ15 NT-S and TDFT-S in mammals are unknown. Methods We fed captive mice (Mus musculus) either (A) a low-fat, high-fiber diet with low, intermediate, or high protein; or (B) a high-fat, low-fiber diet with low or intermediate protein. Mouse muscle and dietary protein were analyzed for bulk tissue δ15 N using elemental analyzer-isotope ratio mass spectrometry (EA-IRMS), and were also hydrolyzed into free AA that were analyzed for δ15 N using gas chromatography-combustion-IRMS. Results As dietary protein increased, Δ15 NConsumer-Diet slightly declined for bulk muscle tissue in both experiments; increased for AA in the low-fat, high-fiber diet (A); and remained the same or decreased for AA in the high-fat, low-fiber diet (B). The effects of dietary protein on Δ15 NT-S and on TDFT-S varied by AA but were consistent between variables. Conclusions Diets were less digestible and included more protein in Experiment A than in Experiment B. As a result, the mice in Experiment A probably oxidized more AA, resulting in greater Δ15 NConsumer-Diet values. However, the similar responses of Δ15 NT-S and of TDFT-S to diet variation suggest that if diet samples are available, Δ15 NT-S accurately tracks trophic position. If diet samples are not available, the patterns presented here provide a basis to interpret Δ15 NT-S values. The trophic-source offset of Pro-Lys did not vary across diets, and therefore may be more reliable for omnivores than other offsets (e.g., Glu-Phe).

Polar bears possess morphologic and physiologic characteristics that reflect their terrestrial lineage as members of the bear family (Ursidae) as well as adaptations to the Arctic marine environment. Among marine mammals, they are the least adapted for aquatic life. They exhibit substantial seasonality in body mass, body condition, and many physiological functions, reflecting the annual cycle of both their Arctic sea ice habitat and the availability of their main prey, ringed seals. This hypercarnivorous diet has likely influenced the polar bear’s craniodental morphology and nutritional physiology. Similar to other marine mammal predators, polar bears exhibit a relatively high resting metabolic rate (RMR) and field metabolic rate (FMR). The polar bear skeleton is well adapted for walking, rather than tree-climbing, and to a lesser degree, for swimming. The large feet provide secure traction on sea ice (aided by sharp claws) and propulsion in the water. Their reproduction, winter hibernation (by pregnant females), and sensory systems resemble those of other bears. Future research should focus on nutrient recycling during fasting, adaptation to a high-fat diet, susceptibility to pathogens, and assessment of the fitness consequences of ongoing sea ice loss and chemical contamination of their habitat.

International audience; Capital breeders accumulate nutrients prior to egg development, then use these stores to support offspring development. In contrast, income breeders rely on local nutrients consumed contemporaneously with offspring development. Understanding such nutrient allocations is critical to assessing life‐history strategies and habitat use. Despite the contrast between these strategies, it remains challenging to trace nutrients from endogenous stores or exogenous food intake into offspring. Here, we tested a new solution to this problem. Using tissue samples collected opportunistically from wild emperor penguins Aptenodytes forsteri, which exemplify capital breeding, we hypothesized that the stable carbon (δ13C) and nitrogen (δ15N) isotope values of individual amino acids (AAs) in endogenous stores (e.g. muscle) and in egg yolk and albumen reflect the nutrient sourcing that distinguishes capital versus income breeding. Unlike other methods, this approach does not require untested assumptions or diet sampling. We found that over half of essential AAs had δ13C values that did not differ between muscle and yolk or albumen, suggesting that most of these AAs were directly routed from muscle into eggs. In contrast, almost all non‐essential AAs differed in δ13C values between muscle and yolk or between muscle and albumen, suggesting de novo synthesis. Over half of AAs that have labile nitrogen atoms (i.e. ‘trophic’ AA) had higher δ15N values in yolk and albumen than in muscle, suggesting that they were transaminated during their routing into egg tissue. This effect was smaller for AAs with less labile nitrogen atoms (i.e. ‘source’ AA). Our results indicate that the δ15N offset between trophic‐source AAs (Δ15Ntrophic‐source) may provide an index of the extent of capital breeding. The value of emperor penguin Δ15NPro‐Phe was higher in yolk and albumen than in muscle, reflecting the mobilization of endogenous stores; in comparison, the value of Δ15NPro‐Phe was similar across muscle and egg tissue in previously published data for income‐breeding herring gulls Larus argentatus smithsonianus. Our results provide a quantitative basis for using AA δ13C and δ15N, and isotopic offsets among AAs (e.g. Δ15NPro‐Phe), to explore the allocation of endogenous versus exogenous nutrients across the capital versus income spectrum of avian reproduction.

The measurement of bulk tissue nitrogen (δ15N) and carbon isotope values (δ13C) chronologically along biologically inert tissues sampled from offspring can provide a longitudinal record of their mothers’ foraging habits. This study tested the important assumption that mother–offspring stable isotope values are positively and linearly correlated. In addition, any change in the mother–offspring bulk tissues and individual amino acids that occurred during gestation was investigated. Whiskers sampled from southern elephant seal pups (Mirounga leonina) and temporally overlapping whiskers from their mothers were analyzed. This included n = 1895 chronologically subsampled whisker segments for bulk tissue δ15N and δ13C in total and n = 20 whisker segments for amino acid δ15N values, sampled from recently weaned pups (n = 17), juvenile southern elephant seals (SES)

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.

Seasonal reductions in food availability may cause animals to catabolize endogenous tissue and the resulting loss of lean mass can hinder their ability to forage and reproduce. While several studies have considered nitrogen isotopes (δ15N) as an indicator of catabolism, relationships between protein intake, body condition, and tissue δ15N have not been assessed simultaneously in controlled conditions. We conducted a feeding experiment on laboratory mice (Mus musculus) to test the effects of low (5%) versus high (30–40%) dietary protein content on lean mass, fat mass, and tissue δ15N. This approach enables the distinction between use of exogenous and endogenous nitrogen, illuminating a framework of protein metabolism and tissue synthesis. As expected, lean mass and body fat were lower in mice fed low-protein diets. Nitrogen isotope discrimination (Δ15N) between blood plasma-diet and liver-diet did not differ between diet treatments. In contrast, Δ15N for hair decreased while Δ15N for muscle and RBC increased in the low-protein treatment. These patterns suggest that animals in negative nitrogen balance catabolize labile endogenous protein (e.g., muscle) to maintain vital tissues (e.g., liver) required to survive. Consequently, muscle and RBC δ15N values appear to be the most useful in assessing the nutritional state of animals. Our combination of direct measurements of body condition with δ15N analysis suggest how nitrogen isotopes can be better used as tracers of catabolic and anabolic activity by demonstrating connections between tissue-specific metabolic processes and Δ15N, thus refining the application of δ15N as a tool for assessing nitrogen balance in wild animals.

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.

Although tissue fatty acid (FA) composition has been linked to whole-animal performance (e.g., aerobic endurance, metabolic rate, postexercise recovery) in a wide range of animal taxa, we do not adequately understand the pace of changes in FA composition and its implications for the ecology of animals. Therefore, we used a C4 to C3 diet shift experiment and compound-specific δ13C analysis to estimate the turnover rates of FAs in the polar and neutral fractions of flight muscle lipids (corresponding to membranes and lipid droplets) of exercised and sedentary zebra finches (Taeniopygia guttata). Turnover was fastest for linoleic acid (LA; 18:2n6) and palmitic acid (PA; 16:0), with 95% replacement times of 10.8-17.7 d in the polar fraction and 17.2-32.8 d in the neutral fraction, but was unexpectedly slow for the long-chain polyunsaturated FAs (LC-PUFAs) arachidonic acid (20:4n6) and docosahexaenoic acid (22:6n3) in the polar fraction, with 95% replacement in 64.9-136.5 d. Polar fraction LA and PA turnover was significantly faster in exercised birds (95% replacement in 8.5-13.3 d). Our results suggest that FA turnover in intramuscular lipid droplets is related to FA tissue concentrations and that turnover does not change in response to exercise. In contrast, we found that muscle membrane FA turnover is likely driven by a combination of selective LC-PUFA retention and consumption of shorter-chain FAs in energy metabolism. The unexpectedly fast turnover of membrane-associated FAs in muscle suggests that songbirds during migration could substantially remodel their membranes within a single migration stopover, and this may have substantial implications for how the FA composition of diet affects energy metabolism of birds during migration.

The measurement of stable isotopes in ‘bulk’ animal and plant tissues (e.g., muscle or leaf) has become an important tool for studies of functional diversity from organismal to continental scales. In consumers, isotope values reflect their diet, trophic position, physiological state, and geographic location. However, interpretation of bulk tissue isotope values can be confounded by variation in primary producer baseline values and by overlapping values among potential food items. To resolve these issues, biologists increasingly use compound-specific isotope analysis (CSIA), in which the isotope values of monomers that constitute a macromolecule (e.g., amino acids in protein) are measured. In this review, we provide the theoretical underpinnings for CSIA, summarize its methodology and recent applications, and identify future research directions. The key principle is that some monomers are reliably routed directly from the diet into animal tissue, whereas others are biochemically transformed during assimilation. As a result, CSIA of consumer tissue simultaneously provides information about an animal’s nutrient sources (e.g., food items or contributions from gut microbes) and its physiology (e.g., nitrogen excretion mode). In combination, these data clarify many of the confounding issues in bulk analysis and enable novel precision for tracing nutrient and energy flow within and among organisms and ecosystems.

Climate change is altering the distribution of some wildlife species while warming temperatures are facilitating the northward expansion of pathogens, potentially increasing disease risk. Melting of Arctic sea ice is increasingly causing polar bears (Ursus maritimus) of the southern Beaufort Sea (SBS) to spend summer on land, where they may encounter novel pathogens. Here, we tested whether SBS polar bears on shore during summer exhibited greater immune system activity than bears remaining on the sea ice. In addition, we tested whether the type of immune response correlated with body condition, because adaptive responses (slowly developing defenses against specific pathogens) often require less energy than innate responses (rapid defenses not based on pathogen identity). After accounting for body condition, we found that polar bears on shore exhibited higher total white blood cell counts, neutrophils, and monocytes than bears on the ice, suggesting more infections. Lymphocytes, eosinophils, basophils, and globulins did not differ. C-reactive protein, an indicator of inflammation, also did not differ between habitats. Body condition was associated with variables indicative of both innate and adaptive immunity, suggesting that neither response was uniquely limited by energy resources. Our data indicate that as more polar bears spend longer periods of time on shore, they may experience more infections. We encourage continued health monitoring of this species and studies of the long-term fitness consequences from disease.

Animal keepers at zoos and wildlife rescue centers often possess in-depth knowledge of the health and behavior of the individuals under their care. While it is often not feasible for keepers to regularly collect behavior data through formal scientific methods, efforts should be made to find alternative means to capture this knowledge. We investigated the use of keeper feedback to study the behavior of sloth bears at the Agra Bear Rescue Facility (ABRF; Agra, India). We prepared a survey with 5 questions focused on behaviors indicative of playfulness, boldness, aggressiveness, and the tendency to express self-directed behaviors (SDB). We asked keepers to rate bears on a Likert scale from 1 (least likely to exhibit a behavior) to 5 (most likely) for 44 adult female bears (5-21 years of age). We validated this method by comparing keeper ratings of SDB with formal behavior observations, finding that time of day had an influence on the accuracy of keeper assessments. We found a significant negative correlation between housing bears in larger groups (>15) and SDB. In addition, we correlated ratings given by keepers for all study behaviors. Social play had significant negative correlation with aggression toward people. There was no correlation between social play and aggression toward other bears, possibly due to the existence of cohesive social groups in group housing or high dimensionality of the data. We found that keeper feedback is an efficient tool to gather behavior data on captive sloth bears and recommend its use in future studies.

Not that unusual after all As polar ice recedes, polar bears are facing a changed habitat with reduced summer foraging opportunities. It has been hypothesized that they might be able to resist summer food shortages by reducing their metabolic needs in a sort of “walking hibernation.” Whiteman et al. monitored energy expenditure in polar bears both on and off the ice and found energy reductions, but that these were more akin to normal mammalian fasting levels. Thus, it appears that polar bears have no energetic protections against reduced summer food supplies and will face increasing starvation threats if summer foraging habitats continue to decline. Science , this issue p. 295

Polar bears experience skeletal muscle atrophy during winter whether they hibernate or not. Bears spending summer on sea ice in the Arctic basin undergo a second period of atrophy caused by fasting, whereas bears with shore access to adequate, alternative food do not. Summer atrophy may influence fitness., When reducing activity and using stored energy during seasonal food shortages, animals risk degradation of skeletal muscles, although some species avoid or minimize the resulting atrophy while experiencing these conditions during hibernation. Polar bears may be food deprived and relatively inactive during winter (when pregnant females hibernate and hunting success declines for other demographic groups) as well as summer (when sea ice retreats from key foraging habitats). We investigated muscle atrophy in samples of biceps femoris collected from free-ranging polar bears in the Southern Beaufort Sea (SBS) throughout their annual cycle. Atrophy was most pronounced in April–May as a result of food deprivation during the previous winter, with muscles exhibiting reduced protein concentration, increased water content, and lower creatine kinase mRNA. These animals increased feeding and activity in spring (when seal prey becomes more available), initiating a period of muscle recovery. During the following ice melt of late summer, ~30% of SBS bears abandon retreating sea ice for land; in August, these ‘shore’ bears exhibited no muscle atrophy, indicating that they had fully recovered from winter food deprivation. These individuals subsequently scavenged whale carcasses deposited by humans and by October, had retained good muscle condition. In contrast, ~70% of SBS bears follow the ice north in late summer, into deep water with less prey. These ‘ice’ bears fast; by October, they exhibited muscle protein loss and rapid changes in myosin heavy-chain isoforms in response to reduced activity. These findings indicate that, unlike other bears during winter hibernation, polar bears without food in summer cannot mitigate atrophy. Consequently, prolonged summer fasting resulting from climate change-induced ice loss creates a risk of greater muscle atrophy and reduced abilities to travel and hunt.

Seasonal reductions in food availability may cause animals to catabolize endogenous tissue and the resulting loss of lean mass can hinder their ability to forage and reproduce. While several studies have considered nitrogen isotopes (δ

As human activities lead to rising greenhouse gas concentrations in Earth's atmosphere, less incoming solar energy is released back into space, causing a net energy gain that increases global temperatures. The consequence of climate change for polar bears can likewise be understood in terms of an energy imbalance. Sea ice melting reduces the opportunities for polar bears to capture seals (see the photo), leaving them at risk of expending more energy in the pursuit of food than they can obtain. The magnitude of this imbalance is determined by their rate of energy use. On page 568 of this issue, Pagano et al. ( 1 ) quantify the energy expense of wild polar bears and show that it is higher than previously estimated.

For many terrestrial organisms in the Northern Hemisphere, winter is a period of resource scarcity and energy deficits, survivable only because a seasonal refugium – the “subnivium” – exists beneath the snow. The warmer and more stable conditions within the subnivium are principally driven by snow duration, density, and depth. In temperate regions, the subnivium is important for the overwintering success of plants and animals, yet winter conditions are changing rapidly worldwide. Throughout the Northern Hemisphere, the impacts of climate change are predicted to be most prominent during the winter months, resulting in a shorter snow season and decreased snow depth. These climatic changes will likely modify the defining qualities of the subnivium, resulting in broad-scale shifts in distributions of species that are dependent on these refugia. Resultant changes to the subnivium, however, will be spatially and temporally variable. We believe that ecologists and managers are overlooking this widespread, crucia...

Animals moving across snow surfaces sink to varying depths, increasing the energetic cost of travel. For ease of movement, animals may follow compacted trails created by sports such as snowmobiling and snowshoeing. We tested the assumption that animals less-adapted to snow travel (i.e. animals with a high footload (body mass/foot surface area)) are more likely to use compacted trails and follow them for greater distances than animals well-adapted to moving on snow. We sampled animal movements on compacted and non-compacted transects in northwestern Wyoming and southeastern Idaho, USA, during two winters. Consistent with our prediction, footload positively influenced the probability of animals following compacted transects, and positively influenced their following distance, although the latter relationship was weaker. Additionally, we found that different snow sports cause similar increases in snow density. We suggest that future researchers experimentally test whether the use of compacted trails by anima...

The incorporation of dietary macronutrients and associated isotopic signatures of carbon (δ13C) and nitrogen (δ15N) into animal tissues is a result of the interaction between growth, nutritional status, and the composition of the diet. In mammalian carnivores incorporation is further complicated by lack of carbohydrates in the diet and allocation of large quantities of dietary macronutrients to fetuses and milk production. In this study, we explored the effects of diet composition, growth, pregnancy, and milk production on isotopic incorporation of 13C and 15N in captive mink (Neovison vison) fed 3 experimental diets (Beef, Fish, and a Mixture of the 2) that differed in lipid and amino acid composition. In nursing kits, growth was the main factor influencing isotopic incorporation rates into muscle. Similarly, in adults, changes in body mass influenced the dynamics of isotopic incorporation in red blood cells, although the rates differed for δ13C and δ15N, as well as among the 3 experimental grou...

Dietary carbon is oxidized and exhaled as CO2, thus δ13Cbreath values can provide information on diet and substrate use for energy. However, physiological phenomena such as fat deposition and fasting can alter values of δ13Cbreath such that interpretation of source contributions may be unclear. Consequently, before application to free-ranging animals, inferences about feeding and nutritional states based on δ13Cbreath should be validated with controlled experiments using captive individuals. Here, we report δ13Cbreath values for 4 captive brown bears (Ursus arctos) under different conditions: the bears were 1st given a diet containing carbohydrate, lipid, and protein; they were then switched to a carbohydrate-free diet consisting of salmon and fish oil; and finally they were placed on a fast leading to winter hibernation. Following the switch to the carbohydrate-free diet, values of δ13Cbreath and δ13Cplasma suggested that although oxidation included a substantial portion of dietary proteins, die...

Demographic structure is central to understanding the dynamics of animal populations. However, determining the age of free-ranging mammals is difficult, and currently impossible when sampling with noninvasive, genetic-based approaches. We present a method to estimate age class by combining measures of telomere lengths with other biologically meaningful covariates in a Bayesian network. We applied this approach to American and Pacific martens (Martes americana and M. caurina) and compared predicted age with that obtained from counts of cementum annuli. Using telomere length and the covariates sex, species, and estimates of population density obtained from commercial trapping records, we assigned martens to juvenile (

Polar bears (Ursus maritimus) prefer to live on Arctic sea ice but may swim between ice floes or between sea ice and land. Although anecdotal observations suggest that polar bears are capable of swimming long distances, no data have been available to describe in detail long distance swimming events or the physiological and reproductive consequences of such behavior. Between an initial capture in late August and a recapture in late October 2008, a radio-collared adult female polar bear in the Beaufort Sea made a continuous swim of 687 km over 9 days and then intermittently swam and walked on the sea ice surface an additional 1,800 km. Measures of movement rate, hourly activity, and subcutaneous and external temperature revealed distinct profiles of swimming and walking. Between captures, this polar bear lost 22% of her body mass and her yearling cub. The extraordinary long distance swimming ability of polar bears, which we confirm here, may help them cope with reduced Arctic sea ice. Our observation, however, indicates that long distance swimming in Arctic waters, and travel over deep water pack ice, may result in high energetic costs and compromise reproductive fitness.

We examined differences between summer and winter in xanthophyll cycle-dependent energy dissipation and leaf antioxidant systems in needles of the overwintering evergreen Taxus x media cv. Tauntonii (Taunton yew) growing in both sun and shade environments in Saint Paul, Minnesota. During the winter, both sun and shade plants exhibited increases in the capacity for, and utilization of, xanthophyll cycle-dependent thermal energy dissipation. Winter needles showed decreases (sun needles) or no change (shade needles) in superoxide dismutase activity (EC 1.15.1.1), no change in ascorbate peroxidase activity (EC 1.11.1.11) and no change (sun needles) or increases (shade needles) in reduced ascorbate levels. Both sun and shade needles showed large increases in glutathione reductase activity (EC 1.6.4.2) and total glutathione levels during the winter, in addition to increases in levels of a-tocopherol. These results suggest an important photoprotective role during the winter for xanthophyll cycle-dependent energy dissipation and for the antioxidants glutathione and a-tocopherol. They suggest a less important photoprotective function of the enzyme-based water–water cycle in winter acclimation in the seasonally very cold environment of Minnesota.

Blood triacylglycerol (TG) and lipoproteins are important variables for evaluating nutritional status of wildlife, but measurements are often expensive and difficult. Performance of a small, portable blood analyzer intended for human medical diagnostics was evaluated in measuring these variables in plasma and serum from free-ranging polar bears ( Ursus maritimus), which are experiencing nutritional stress related to sea ice loss. The analyzer accurately tracked changes in concentration of total cholesterol (Ctotal), cholesterol associated with high-density lipoprotein (CHDL), and TG during a validation protocol of diluting samples and spiking them with exogenous cholesterol and glycerol. Values of Ctotal and TG agreed well with values obtained by other methods (ultracentrifugation followed by colorimetric assays); agreement was variable for values of cholesterol associated with specific lipoproteins. Similar to a study of captive polar bears, ultracentrifugation methods revealed greater TG in very low-density lipoproteins than in low-density lipoprotein, which is unusual and merits additional study.