Arctic ground squirrels overwintering in northern Alaska experience average soil temperature of -10 degrees C. To examine energetic costs of arousing from hibernation under arctic compared to temperate conditions, captive ground squirrels were maintained in ambient temperatures (T(a)) of 2, -5 and -12 degrees C. Rates of oxygen consumption and carbon dioxide production were used to estimate metabolic rate and fuel use during the three phases of arousal episodes: rewarming, euthermia, and recooling. Respiratory quotient comparisons suggest exclusive use of lipid during rewarming and mixed fuel use during euthermia. Animals rewarming from torpor at T(a) -12 degrees C took longer, consumed more oxygen, and attained higher peak rates of oxygen consumption when compared to 2 degrees C. T(a) had no significant effect on cost or duration of the euthermic phase. Animals recooled faster at -12 degrees C than at 2 degrees C, but total oxygen consumption was not different. T(a) had no significant effect on the total cost of arousal episodes when all three phases are included. Arousal episodes account for 86% of estimated costs of a complete hibernation cycle including torpor when at 2 degrees C and only 23% at -12 degrees C. Thus, due to the higher costs of steady-state metabolism during torpor, proportional metabolic costs of arousal episodes at T(a) characteristic of the Arctic are diminished compared to relative costs of arousals in more temperate conditions.

Among pinnipeds, harbor seals Phoca vitulina have the broadest distribution (a 34° to 50° range in latitudes in the Pacific and Atlantic regions, respectively) and are found in a diversity of habitats. Harbor seals in Alaska, USA, similar to Arctic pinnipeds in many respects, rely upon glacial ice for pupping, mating, and molting. Just as climate change affects Arctic sea ice, tidewater glaciers are rapidly retreating in Alaska, reducing ice availability for harbor seals. An increased understanding of glacial vs. terrestrial harbor seals may reveal information important to conservation of harbor seals and Arctic pinnipeds, as effects of climate change continue. We compared foraging distances, activity budgets, diet, and body condition for seals captured at glacial and terrestrial sites in Glacier Bay, Alaska. Foraging strategies and activity budgets of seals using glacial ice differed substantially from seals using terrestrial sites. Glacial seals traveled significantly farther to forage (≥40 vs. < 5 km) and spent more time hauled out than terrestrial seals (26 vs. 11 to 16%). Diets of glacial seals were higher in pelagic fishes compared to diets of terrestrial seals that foraged primarily on intertidal/demersal fishes. Body condition of seals was similar between habitats (p ≥ 0.09) and suggests that costs of longer foraging trips for glacial seals may be offset by obtaining higher quality diets of pelagic fishes, which may allow seals to spend more time hauled out. During the brief lactation period, more time hauled out could result in more time available for the transfer of energy from adult females to dependent offspring. KEY WORDS: Foraging behaviour · Activity budget · Body condition · Habitat use · Stable isotopes · Time-depth recorder · Phocid · Pinniped Resale or republication not permitted without written consent of the publisherMar Ecol Prog Ser 429: [277][278][279][280][281][282][283][284][285][286][287][288][289][290] 2011 numbers of pupping harbor seals (Calambokidis et al. 1987), suggesting that these sites provide important pupping and breeding habitats. Steep declines in harbor seal numbers have occurred at several glacial ice sites in Alaska (Hoover-Miller 1994, 2011, Mathews & Pendleton 2006, Womble et al. 2010, highlighting the need for a better understanding of the basic ecology of seals using glacial habitat and the potential benefits and costs of using glacial ice to pup, breed, or molt.Glacier Bay National Park seasonally hosts one of the largest aggregates of harbor seals in Alaska. Seals use both terrestrial substrate and glacial ice within the Bay, which provides a unique opportunity to understand the role of different habitats in influencing seal behavior within a relatively small geographic region. Also of interest is whether a comparison of seals in ice and terrestrial habitats can be used as a corollary to the situation in the Arctic as it relates to diminishing sea ice (Rodrigues 2009, Polyak et al. 2010 and Arctic phocids that rely upon ice substrate for resting, fora...

Laboratory experiments have demonstrated that the amount of polyunsaturated fatty acids (PUFAs) in the diet before hibernation influences patterns of mammalian torpor. The hibernation ability of ground squirrels is greatest (longest torpor bouts, greatest number of animals entering torpor) when the PUFA content of their fall diets is 33-74 mg/g, under laboratory conditions. The extent to which natural fall diets both (a) vary in PUFA content and (b) influence the torpor patterns of free-ranging populations of hibernating mammals is unknown, however. We conducted a 3-yr study on the diet PUFA contents and subsequent hibernation patterns of free-ranging arctic ground squirrels (Spermophilus parryii) in the Brooks Range of Alaska. We found that the PUFA contents of fall diets varied more than threefold among individuals. Our study also revealed that arctic ground squirrels that consumed a moderate-PUFA (33-74 mg/g) diet had (a) longer torpor bouts, (b) fewer arousals from torpor, (c) shorter arousal periods, (d) more days in torpor, and (e) greater probability of persisting in the population than those that consumed a high-PUFA (>74 mg/g) diet during the fall. No animals were demonstrated to have consumed a diet representing low-PUFA (<33 mg/g) values. Our study is therefore the first to demonstrate that estimated dietary PUFA levels of a free-ranging hibernator influence subsequent torpor patterns.

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Alaskan seals are found in remote and sometimes inaccessible locations, making it difficult to collect time‐series information. This study explores a novel method to examine temporal changes in diet and physiological status of ringed (Pusa hispida), spotted (Phoca largha), and harbor (Phoca vitulina) seals using cortisol concentrations and δ15N and δ13C stable isotopes (SIs) measured in serial sections of whiskers. As whiskers grow, whisker tissue is deposited sequentially making these measurements temporally aligned. Whisker cortisol presented in a distinct pattern with elevated concentrations at the root section followed by a curvilinear decline moving toward the tip of most whiskers. Comparing SIs at the root to the rest of the whiskers, δ13C values were slightly lower in ringed and harbor seal whiskers and δ15N values were slightly higher in harbor seal whiskers. The data were modeled controlling for the observed trends in cortisol concentrations and further associations between cortisol concentrations and SIs were detected in spotted and harbor seal whiskers. Additional research examining the source and stability of whisker cortisol is warranted. However, the methods presented here demonstrate that whiskers could prove valuable to gather long‐term and naturally aligned dietary and physiological information.

Ringed (Pusa hispida) and bearded seals (Erignathus barbatus) inhabit vast and often remote areas in the Arctic, making it difficult to obtain long-term physiological information concerning health and reproduction. These seals are experiencing climate-driven changes in their habitat that could result in physiological stress. Chronic physiological stress can lead to immunosuppression, decreased reproduction and decreased growth. Recently, keratin has become a popular matrix to measure steroid hormones, such as stress-related cortisol and reproduction-related progesterone. We developed and validated methods to extract cortisol and progesterone from the claws of adult female ringed (n = 20) and bearded (n = 3) seals using enzyme immunosorbent assays. As ringed and bearded seal claws grow, a pair of dark- and light-colored bands of keratin is deposited annually providing a guide for sampling. Two processing methods were evaluated, removal of claw material with a grinding bit or grinding followed by mechanical pulverization (102 paired samples from six claws, two each from three seals). Adding the mechanical pulverization step resulted in a 1.5-fold increase in hormone extraction. Progesterone from the proximal claw band was evaluated to biologically validate claw material as a measure of pregnancy in ringed seals (n = 14). Claws from pregnant seals had significantly higher claw progesterone concentrations than from non-pregnant seals. This suggests that the elevated progesterone associated with gestation was reflected in the claws, and that the most proximal claw band was indicative of pregnancy status at time of death. Thus, although the sample size was low and the collection dates unbalanced, this study demonstrates the potential to use claws to examine an extended time series (up to 12 yrs) of cortisol and progesterone concentrations in ringed and bearded seal claws.

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Rapid climate warming is decreasing sea ice thickness, extent and duration. Marine mammals such as bearded (Erignathus barbatus) and ringed (Pusa hispida) seals, which use sea ice for pupping, molting and resting, may be negatively affected. Claws from bearded and ringed seals store up to 14 and 12 years of sequential analyte data, respectively. These data can be used to compare reproduction, stress and diet across decades. In this study, we compare progesterone, cortisol and carbon and nitrogen stable isotopes in female bearded and ringed seals during 1953–1968 (pre-1968, a period prior to sea ice decline) to 1998–2014 (post-1998, a period during sea ice decline). When comparing these periods, bearded seals had statistically higher cortisol concentrations post-1998, and for both species δ13C was more negative post-1998, while progesterone and δ15N did not change. There was a positive relationship between progesterone and cortisol Z-scores for both species, except for ringed seals post-1998. There was a negative relationship between cortisol Z-scores and δ13C for bearded seals evident in post-1998 indicating that higher cortisol Z-scores are associated with more negative δ13C in bearded seals in recent years. This negative relationship between cortisol and δ13C in bearded seals suggests a shift to higher prey diversity, possibly due to changes in sea ice in the Pacific Arctic evident post 1998. Progesterone Z-scores corresponded to expected differences among non-pregnant, unimplanted, implanted and post-partum individuals. Using these data, pregnancy history was determined for reproductive years for each individual female sampled, which could allow for yearly pregnancy rates to be calculated given a large enough representative sample of the population. These results combine decades of observational studies with hormones and stable isotopes to infer changes in reproduction, stress and diet, as well as the connection between these life history parameters.