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.

A novel method to measure steroid hormone concentrations in walrus bone from archaeological, historical, and modern time periods using liquid chromatography/tandem mass spectrometry.

RATIONALE: A liquid chromatography/tandem mass spectrometry (LC/MS/MS) method was validated and utilized to measure and analyze four steroid hormones related to stress and reproduction in individual samples from a novel tissue, Pacific walrus (Odobenus rosmarus divergens, herein walrus) bone. This method determines steroid hormone concentrations in the remote walrus population over millennia from archaeological (>200 bp), historical (200-20 bp), and modern (2014-2016) time periods. METHODS: Lipids were extracted from walrus bone collected from these periods using methanol before LC/MS/MS analysis. Isotopically labeled internal standards for each target hormone were added to every sample. Analytical and physiological validations were performed. Additionally, a tissue comparison was done among paired walrus bone, serum, and blubber samples. A rapid resolution liquid chromatography system coupled to a QqQ mass spectrometer was used to analyze all samples after derivatization for progesterone, testosterone, cortisol, and estradiol concentrations. Multiple reaction monitoring was used for MS analysis and data were acquired in positive electrospray ionization mode. RESULTS: Progesterone, testosterone, cortisol, and estradiol were linear along their respective standard calibration curves based on their R2 values (all > 0.99). Accuracy ranged from 93-111% for all hormones. The recovery of extraction, recovery of hormones without matrix effect, was 92-101%. The overall process efficiency of our method for measuring hormones in walrus bone was 93-112%. Progesterone and testosterone concentrations were not affected by reproductive status among adult females and males, respectively. However, estradiol was different among pregnant and non-pregnant adult females. Overall, steroid hormones reflect a long-term reservoir in cortical bone. This method was also successfully applied to walrus bone as old as 3585 bp. CONCLUSIONS: LC/MS/MS analysis of bone tissue (0.2-0.3 g) provides stress and reproductive data from elusive walruses that were alive thousands of years ago. Based on physiological validations, tissue comparison, and published literature, steroid hormone concentrations measured in walrus cortical bone could represent an accumulated average around a 10-20-year time span. By investigating how stress and reproductive physiology may have changed over the past ~3000 years based on bone steroid hormone concentrations, this method will help answer how physiologically resilient walruses are to climate change in the Arctic.

Organochlorine pesticide accumulation in seabirds and marine mammals from the Northwest Pacific.

Bioaccumulation of organochlorine pesticides by marine organisms is one of the methods of environmental quality. Concentrations of organochlorine pesticides (HCH isomers (α-, ß-, γ-HCH), p,p'-DDT and its metabolites (p,p'-DDD, p,p'-DDE)) in samples of seabirds (Northern fulmar Fulmarus glacialis, Crested auklet Aethia cristatella, Auklet-crumb Aethia pusilla, Pacific gull Larus schistisagus, and Gray petrel Oceanodroma furcate) and marine mammals (Gray whales Eschrichtius robustus and Pacific walrus Odobenus rosmarus divergens) were measured by a GC-MS. The total concentration of OCPs in mammals was higher than in seabirds. Environmental biomonitoring with the use of seabirds and marine mammals, as a long-lived species, is used for global monitoring, since bioaccumulation in these organisms occurs throughout life. The environmental assessment of the Northwest Pacific marine ecosystems revealed that the levels of OCP contamination in this area are similar to the other subarctic regions of the World Ocean.

Sex-Specific Energetics of Pacific Walruses (Odobenus rosmarus divergens) during the Nursing Interval.

Habitat use and activity patterns of Pacific walruses (Odobenus rosmarus divergens) have changed with climate-induced reductions in sea ice. Increases in the time active in water could result in negative energy balance, precluding females from sustaining lactation, which could impact population demographics. Little is known about lactation costs in walruses. We examined the energetics of 0-2-yr-old walrus calves by using Bayesian hierarchical models based on longitudinal husbandry records of growth (n = 6 females and 7 males) and caloric intake (n = 5 females and 6 males) as a proxy for maternal lactation costs. Males and females had similar growth patterns; mean mass increased from 68 kg at birth to 301 kg by 2 yr. Females had a 2,000 kcal kg(-1) higher mass storage (growth) cost than males; females typically synthesize and deposit greater amounts of adipose, which is more energy dense than lean tissue. In contrast, males had higher metabolic (basal and activity) costs, ranging from 600 to 1,800 kcal d(-1) greater than similarly sized females; males are typically leaner, and muscle is more metabolically active than adipose. Yet total daily energy requirements (storage plus metabolic components) were similar across sexes, summing to approximately 190,000 kcal over the first month postpartum. Based on these estimates and assuming that 8,103 kcal is recovered from 1 kg of mass loss in adult female walruses, suckling calves could deplete 23 kg of their mother's body mass over the first month after parturition if none of the lactation costs is met through ingested prey.

Rapid maturation of the muscle biochemistry that supports diving in Pacific walruses (Odobenus rosmarus divergens).

Physiological constraints dictate animals' ability to exploit habitats. For marine mammals, it is important to quantify physiological limits that influence diving and their ability to alter foraging behaviors. We characterized age-specific dive limits of walruses by measuring anaerobic (acid-buffering capacity) and aerobic (myoglobin content) capacities of the muscles that power hind (longissimus dorsi) and fore (supraspinatus) flipper propulsion. Mean buffering capacities were similar across muscles and age classes (a fetus, five neonatal calves, a 3 month old and 20 adults), ranging from 41.31 to 54.14 slykes and 42.00 to 46.93 slykes in the longissimus and supraspinatus, respectively. Mean myoglobin in the fetus and neonatal calves fell within a narrow range (longissimus: 0.92-1.68 g 100 g(-1) wet muscle mass; supraspinatus: 0.88-1.64 g 100 g(-1) wet muscle mass). By 3 months post-partum, myoglobin in the longissimus increased by 79%, but levels in the supraspinatus remained unaltered. From 3 months post-partum to adulthood, myoglobin increased by an additional 26% in the longissimus and increased by 126% in the supraspinatus; myoglobin remained greater in the longissimus compared with the supraspinatus. Walruses are unique among marine mammals because they are born with a mature muscle acid-buffering capacity and attain mature myoglobin content early in life. Despite rapid physiological development, small body size limits the diving capacity of immature walruses and extreme sexual dimorphism reduces the diving capacity of adult females compared with adult males. Thus, free-ranging immature walruses likely exhibit the shortest foraging dives while adult males are capable of the longest foraging dives.

Bioaccumulation of persistent organochlorine pesticides (OCPs) by gray whale and Pacific walrus from the western part of the Bering Sea.

The feeding habits of a gray whale (Eschrichtius robustus) and a Pacific walrus (Odobenus rosmarus divergens), caught from the western Bering Sea in the summers of 2010 and 2011, have been studied, and concentration of persistent organochlorine pesticides (OCPs) in their organs determined. The total OCP concentration (∑HCH+∑DDT) in muscles and liver of the gray whales varies from 297 to 3581 and from 769 to 13,808 ng/g lipids, respectively. The total OCP concentration (∑HCH+∑DDT) in muscles and liver of the Pacific walruses varies from 197 to 5659 and from 4856 to 90,263 ng/g lipids, respectively. The specifics of diet as a source of pesticide accumulation in these two marine mammal species are discussed.

Temporal trends of mercury in marine biota of west and northwest Greenland.

Temporal trends in mercury concentrations ([Hg]) during the last two to three decades were determined in liver of shorthorn sculpin, ringed seal and Atlantic walrus from northwest Greenland (NWG, 77 degrees N) and in liver of shorthorn sculpin and ringed seal from central west Greenland (CWG, 69 degrees N) during the last decade. Stable-nitrogen (delta(15)N) and carbon (delta(13)C) isotope values were determined in muscle of ringed seals to provide insight into potential trophic level changes through time. Log-linear regressions on annual median [Hg] did not reveal any temporal trend in shorthorn sculpin from CWG and NWG and walrus from NWG. In ringed seals from NWG, an increase in [Hg] of 7.8% per year was observed. When based on delta(15)N-adjusted [Hg] this rate increased to 8.5% but was still non-significant. In ringed seal from CWG no trend was found in [Hg] during the period 1994-2004. However, during the last part of the period (1999-2004) the [Hg] increased significantly. Including tissue delta(15)N values as a covariate had a marked effect on these results. The annual changes in delta(15)N-adjusted [Hg] was estimated to -5.0% for the whole period and 2.2% during the last 5 years compared to -1.3% and 12.4%, respectively, for the non-adjusted [Hg].

Congener-specific accumulation and patterns of chlorinated and brominated contaminants in adult male walruses from Svalbard, Norway: indications for individual-specific prey selection.

Blubber samples from 17 adult, male walruses were sampled in eastern Svalbard and analyzed for chlorinated and brominated contaminants. A wide range of contaminants were detected, including PCBs (mean 2000; 95% range 1165-4005 ng/g lipid), DDE (mean 100: 95% range 50-310) ng/g lipid), chlordanes (mean 2500; 95% range 1347-5009) ng/g lipid, toxaphenes (mean 80; 95% range 51-132 ng/g lipid) and polybrominated diphenyl ethers (PBDEs) (mean 15 ng/g; 95% range 9-27 ng/g lipid). PCB and DDE levels were substantially lower than those of animals sampled 10 year earlier in this area, confirming a decreasing trend for these compounds in the Arctic. However, compared to other recently sampled marine mammals from Svalbard, walruses showed relatively high PCB and chlordane levels although they had lower levels of DDE, toxaphenes, and PBDEs, possibly due to species- and location-specific differences in exposure and metabolism. The range in contaminant levels found within the sample group was vast, despite the fact that the animals investigated were all adult males from the same location. The PCB pattern in highly contaminated animals was different from that in animals with low levels of contamination, with relatively more persistent PCBs in the highly contaminated group. This suggests that the more contaminated animals were feeding at higher trophic levels; possibly targeting seals in addition to mollusks as their prey. This suggestion was reinforced by the fatty acid profiles of the inner blubber layer of walruses with low versus high contaminant levels, which suggested different diets for the two groups.

Organochlorine compounds and aliphatic hydrocarbons in Pacific walrus blubber.

Blubber samples were collected from 8 male and 19 female Pacific walrus (Odobenus rosmarus divergens) taken during a 1991 joint USA/USSR cruise traveling widely through the Bering Sea. Dieldrin was found at a level similar to that reported 10 years earlier; oxychlordane was found at a slightly higher concentration than reported previously (Taylor et aL, 1989). Heptachlor epoxide was detected for the first time and found at a low concentration. An initial testing for alpha-, beta- and gamma-HCH detected concentrations similar to those in other Bering Sea pinnipeds. Mean summation of PCB was 0.45 microg g(-1) wet weight in males and 0.16 microg g(-1) in females; only one sample was > 1 microg g(-1). Traces of aliphatic hydrocarbons were detected in all sampled animals, only pristane (x = 0.48 microg g(-1)) was found in concentrations > 1 microg g(-1). Small sample sizes, a lack of samples from immature animals, and uniformly low concentrations of contaminants precluded meaningful analysis of age-related effects and regional differences.