Ursids evolved dietary diversity without major alterations in metabolic rates.

The diets of the eight species of ursids range from carnivory (e.g., polar bears, Ursus maritimus) to insectivory (e.g., sloth bears, Melursus ursinus), omnivory (e.g., brown bears, U. arctos), and herbivory (e.g., giant pandas, Ailuropoda melanoleuca). Dietary energy availability ranges from the high-fat, highly digestible, calorically dense diet of polar bears (~ 6.4 kcal digestible energy/g fresh weight) to the high-fiber, poorly digestible, calorically restricted diet (~ 0.7) of giant pandas. Thus, ursids provide the opportunity to examine the extent to which dietary energy drives evolution of energy metabolism in a closely related group of animals. We measured the daily energy expenditure (DEE) of captive brown bears in a relatively large, zoo-type enclosure and compared those values to previously published results on captive brown bears, captive and free-ranging polar bears, and captive and free-ranging giant pandas. We found that all three species have similar mass-specific DEE when travel distances and energy intake are normalized even though their diets differ dramatically and phylogenetic lineages are separated by millions of years. For giant pandas, the ability to engage in low-cost stationary foraging relative to more wide-ranging bears likely provided the necessary energy savings to become bamboo specialists without greatly altering their metabolic rate.

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.

High-energy, high-fat lifestyle challenges an Arctic apex predator, the polar bear.

Regional declines in polar bear (Ursus maritimus) populations have been attributed to changing sea ice conditions, but with limited information on the causative mechanisms. By simultaneously measuring field metabolic rates, daily activity patterns, body condition, and foraging success of polar bears moving on the spring sea ice, we found that high metabolic rates (1.6 times greater than previously assumed) coupled with low intake of fat-rich marine mammal prey resulted in an energy deficit for more than half of the bears examined. Activity and movement on the sea ice strongly influenced metabolic demands. Consequently, increases in mobility resulting from ongoing and forecasted declines in and fragmentation of sea ice are likely to increase energy demands and may be an important factor explaining observed declines in body condition and survival.

Isotopic Incorporation and the Effects of Fasting and Dietary Lipid Content on Isotopic Discrimination in Large Carnivorous Mammals.

There has been considerable emphasis on understanding isotopic discrimination for diet estimation in omnivores. However, discrimination may differ for carnivores, particularly species that consume lipid-rich diets. Here, we examined the potential implications of several factors when using stable isotopes to estimate the diets of bears, which can consume lipid-rich diets and, alternatively, fast for weeks to months. We conducted feeding trials with captive brown bears (Ursus arctos) and polar bears (Ursus maritimus). As dietary lipid content increased to ∼90%, we observed increasing differences between blood plasma and diets that had not been lipid extracted (∆(13)Ctissue-bulk diet) and slightly decreasing differences between plasma δ(13)C and lipid-extracted diet. Plasma Δ(15)Ntissue-bulk diet increased with increasing protein content for the four polar bears in this study and data for other mammals from previous studies that were fed purely carnivorous diets. Four adult and four yearling brown bears that fasted 120 d had plasma δ(15)N values that changed by <±2‰. Fasting bears exhibited no trend in plasma δ(13)C. Isotopic incorporation in red blood cells and whole blood was ≥6 mo in subadult and adult bears, which is considerably longer than previously measured in younger and smaller black bears (Ursus americanus). Our results suggest that short-term fasting in carnivores has minimal effects on δ(13)C and δ(15)N discrimination between predators and their prey but that dietary lipid content is an important factor directly affecting δ(13)C discrimination and indirectly affecting δ(15)N discrimination via the inverse relationship with dietary protein content.