Indication of two Pacific walrus stocks from whole tooth elemental analysis

The Pacific walrus (Odobenus rosmarus divergens) is considered to be a single panmictic population for management purposes. However, studies on population structuring in this species are limited; in part, because portions of the population’s range are often inaccessible. Therefore, alternative and complementary methods for investigating stock structure in the Pacific walrus are of particular interest. We used measures of elemental concentrations in whole tooth sections from ICP-MS in a discriminant analysis to investigate evidence of stock separation between walruses from two of three known breeding areas (S.E. Bering, St Lawrence, and Anadyr Gulf). Elemental compositions of teeth from female and male walruses from the S.E. Bering and St Lawrence breeding areas were significantly different, providing evidence of separate stocks. We also obtained insights into the potential relation of walruses from non-breeding areas to walruses from these breeding groups based on similarities in their dental elemental profiles.     

Sequence variation at the major histocompatibility complex locus DQ beta in beluga whales (Delphinapterus leucas) [published erratum appears in Mol Biol Evol 1995 Nov;12(6):1174]

Genetic variation at the Major Histocompatibility Complex locus DQ beta was analyzed in 233 beluga whales (Delphinapterus leucas) from seven populations: St. Lawrence Estuary, eastern Beaufort Sea, eastern Chukchi Sea, western Hudson Bay, eastern Hudson Bay, southeastern Baffin Island, and High Arctic and in 12 narwhals (Monodon monoceros) sympatric with the High Arctic beluga population. Variation was assessed by amplification of the exon coding for the peptide binding region via the polymerase chain reaction, followed by either cloning and DNA sequencing or single-stranded conformation polymorphism analysis. Five alleles were found across the beluga populations and one in the narwhal. Pairwise comparisons of these alleles showed a 5:1 ratio of nonsynonymous to synonymous substitutions per site leading to eight amino acid differences, five of which were nonconservative substitutions, centered around positions previously shown to be important for peptide binding. Although the amount of allelic variation is low when compared with terrestrial mammals, the nature of the substitutions in the peptide binding sites indicates an important role for the DQ beta locus in the cellular immune response of beluga whales. Comparisons of allele frequencies among populations show the High Arctic population to be different (P < or = .005) from the other beluga populations surveyed. In these other populations an allele, Dele-DQ beta*0101-2, was found in 98% of the animals, while in the High Arctic it was found in only 52% of the animals. Two other alleles were found at high frequencies in the High Arctic population, one being very similar to the single allele found in narwhal.      

Population structure and gene flow of the Atlantic walrus (Odobenus rosmarus rosmarus) in the eastern Atlantic Arctic based on mitochondrial DNA and microsatellite variation

The population structure of the Atlantic walrus, Odobenus rosmarus rosmarus , was studied using 11 polymorphic microsatellites and restriction fragment length polymorphism detected in the NADH-dehydrogenase ND1, ND2 and ND3/4 segments in mtDNA. A total of 105 walrus samples were analysed from northwest (NW) Greenland, east (E) Greenland, Svalbard and Franz Joseph Land. Two of the 10 haplotypes detected in the four samples were diagnostic for the NW Greenland sample, which implied that the group of walruses in this area is evolutionary distinct from walruses in the other three areas. One individual sampled in E Greenland exhibited a Pacific haplotype, which proved a connection between the Pacific walrus and walruses in eastern Greenland. The Franz Joseph Land, Svalbard and E Greenland samples shared the most common haplotype, indicating very little differentiation at the mtDNA level. Gene flow ( Nm ) estimates among the four areas indicated a very restricted exchange of female genes between NW Greenland and the more eastern Atlantic Arctic samples, and a closer relationship between the three samples composing the eastern Atlantic Arctic. The genetic variation at 11 polymorphic microsatellite loci grouped individuals into three populations, NW Greenland, E Greenland and a common Franz Joseph Land–Svalbard population, which were connected by moderate gene flow.     

Towards a Better Understanding of the Effects of UV on Atlantic Walruses,Odobenus rosmarus rosmarus: A Study Combining Histological Data with Local Ecological Knowledge

Walruses, Odobenus rosmarus, play a key role in the Arctic ecosystem, including northern Indigenous communities, which are reliant upon walruses for aspects of their diet and culture. However, walruses face varied environmental threats including rising sea-water temperatures and decreasing ice cover. An underappreciated threat may be the large amount of solar ultraviolet radiation (UV) that continues to reach the Arctic as a result of ozone loss. UV has been shown to negatively affect whales. Like whales, walrus skin is unprotected by fur, but in contrast, walruses spend long periods of time hauled-out on land. In this study, we combined the results of histological analyses of skin sections from five Atlantic walruses, Odobenus rosmarus rosmarus, collected in Nunavik (Northern Quebec, Canada) with qualitative data obtained through the interviews of 33 local walrus hunters and Inuit Elders. Histological analyses allowed us to explore UV-induced cellular lesions and interviews with experienced walrus hunters and Elders helped us to study the incidences and temporal changes of UV-induced gross lesions in walruses. At the microscopic scale, we detected a range of skin abnormalities consistent with UV damage. However, currently such UV effects do not seem to be widely observed at the whole-animal level (i.e., absence of skin blistering, erythema, eye cataract) by individuals interviewed. Although walruses may experience skin damage under normal everyday UV exposure, the long-term data from local walrus hunters and Inuit Elders did not report a relation between the increased sun radiation secondary to ozone loss and walrus health.     

GROWTH AND SEXUAL DIMORPHISM OF ATLANTIC WALRUSES (ODOBENUS ROSMARUS ROSMARUS) IN FOXE BASIN, NORTHWEST TERRITORIES, CANADA

Growth of Atlantic walruses ( Odobenus rosmarus rosmarus ) was investigated using morphological data collected in association with Inuit subsistence walrus hunts. Four growth models were examined. The growth parameters of a constrained Richards model were used to quantify growth and to test for sexual dimorphism. The asymptotic length of male walruses (315.2 cm ± 3.8 (SE), n = 103) was significantly larger ( t = 7.21, df = 191, P < 0.05) than the asymptotic length of females (276.6 cm ± 3.4, n = 90). Sexual size dimorphism in adults was due to a longer growth period and a faster growth rate in males. The predictive equation relating mass ( M , kg) to standard length ( SL , cm) was: Log₁₀ M = -3.74 + 2.68(Log₁₀ SL ), n = 25, r ²= 0.98. There were no significant differences in the size of male walruses from Foxe Basin collected in the 1950s and this study. There were too few data to compare females. There were no significant differences in size between walruses sampled in Greenland and Foxe Basin in the 1980s and 1990s. Foxe Basin walruses were significantly larger than walruses sampled in northern Hudson Bay in the 1950s. Female Atlantic walruses sampled in Foxe Basin were larger than female Pacific walruses ( Odobenus rosmarus divergens ) sampled in Alaska.     

Serologic evidence of influenza A infection in marine mammals of arctic Canada.

A serologic survey of influenza A antibodies was undertaken on 1,611 blood samples from five species of marine mammals collected from Arctic Canada from 1984-98. Sampling was done in 24 locations throughout the Canadian Arctic encompassing Sachs Harbor (72 degrees N, 125 degrees W), Northwest Territories in the west to Loks Land (63 degrees N, 64 degrees W), Nunavut in the east, to Eureka (80 degrees N, 86 degrees W), Nunavut in the north to Sanikiluaq (56 degrees N, 79 degrees W), Nunavut in the south. A competitive ELISA using a monoclonal antibody (Mab) against influenza A nucleoprotein (NP) was used. Five of 418 (1.2%) belugas (Delphinapterus leucas) and 23 of 903 (2.5%) ringed seals (Phoca hispida) were serologically positive. None of the 210 walruses (Odobenus rosmarus rosmarus), 76 narwhals (Monodon monoceros) and four bowhead whales (Balaena mysticetus) had detectable antibodies to influenza A. Positive belugas were identified from communities on southeast Baffin Island while positive ringed seals came from communities in the eastern, western and high Arctic. Virus isolation attempts on lung tissue from a seropositive beluga were unsuccessful. We believe that influenza A infection in marine mammals is sporadic, the infection is probably self-limiting, and it may not be able to be maintained in these animals. Although the predominant hemagglutinin (H) type was not determined and therefore the pathogenicity of the strains to humans is unknown, the hunting and consumption of marine mammals by the Inuit, may put them at risk for influenza A infection.     

Migration of Walruses (Odobenus rosmarus) in the Svalbard and Franz Josef Land area

Thirty-four walruses ( Odobenus rosmarus ) were fitted with satellite transmitters (PTTs) from 1990 to 1993 in order to study the distribution of the population in the Svalbard area. Twenty-eight were caught at Svalbard and six at Franz Josef Land. All were males except one female caught at Franz Josef Land. At Svalbard, one walrus was caught on the west coast of Spitsbergen, while the others were caught at southern Edgeøya. All walruses were caught in the period from mid-July to early September. The PTTs provided information on location for periods ranging from 0 to 212 days. The results of the satellite trackings show that there is a migration of male walruses between most of the walrus areas at Svalbard and Franz Josef Land. In particular, it seems that migration of males from southern Edgeøya to Kvitøya, Viktoria Island, and Franz Josef Land is common. The walruses winter in the southern parts of Svalbard, as well as within the winter pack-ice of north-eastern Svalbard, which contains numerous open leads. The only walrus at Franz Josef Land that was followed to mid-winter stayed in the area and therefore supports the view that walruses also winter in that area. It is assumed that the majority of walruses at Svalbard are males from one common Svalbard-Franz Josef Land stock. The walrus in the Svalbard-Franz Josef Land area today belong to a recovering population. Their current distribution and behaviour may therefore differ from that found in Svalbard in former times.     

Pacific Walrus (Odobenus rosmarus divergens) Resource Selection in the Northern Bering Sea

The Pacific walrus is a large benthivore with an annual range extending across the continental shelves of the Bering and Chukchi Seas. We used a discrete choice model to estimate site selection by adult radio-tagged walruses relative to the availability of the caloric biomass of benthic infauna and sea ice concentration in a prominent walrus wintering area in the northern Bering Sea (St. Lawrence Island polynya) in 2006, 2008, and 2009. At least 60% of the total caloric biomass of dominant macroinfauna in the study area was composed of members of the bivalve families Nuculidae, Tellinidae, and Nuculanidae. Model estimates indicated walrus site selection was related most strongly to tellinid bivalve caloric biomass distribution and that walruses selected lower ice concentrations from the mostly high ice concentrations that were available to them (quartiles: 76%, 93%, and 99%). Areas with high average predicted walrus site selection generally coincided with areas of high organic carbon input identified in other studies. Projected decreases in sea ice in the St. Lawrence Island polynya and the potential for a concomitant decline of bivalves in the region could result in a northward shift in the wintering grounds of walruses in the northern Bering Sea.     

Giardiasis in pinnipeds from eastern Canada

Cysts of Giardia sp. were detected in feces from the rectum of 20 of 74 pinnipeds examined from the eastern coast of Canada in 1997 and 1998 using a monoclonal antibody technique. Infected pinnipeds included 15 adult harp seals (Phoca groenlandica), four adult grey seals (Halichoerus grypus), and one juvenile harbor seal (Phoca vitulina). Cysts were not detected in 15 seal pups <1-yr-old. The highest prevalence (50%) occurred in adult harp seals collected near the Magdalen Islands in the Gulf of St. Lawrence. The overall prevalence of Giardia sp. in grey and harbor seals, excluding pups, from the Gulf and St. Lawrence estuary was 23%. Feces from 11 beluga (Delphinapterus leucas) and one northern bottle-nosed whale (Hyperoodon ampullatus) stranded in the St. Lawrence estuary were negative for Giardia sp. cysts. The significance of Giardia sp. in marine mammals, shown here for the first time in eastern coastal Canada, is unknown.     

Population structure of North American beluga whales (Delphinapterus leucas) based on nuclear DNA microsatellite variation and contrasted with the population structure revealed by mitochondrial DNA variation

Beluga whales ( Delphinapterus leucas ) in North American waters migrate seasonally between wintering areas in broken pack ice and summering locations in estuaries and other open water areas in the Arctic and sub-Arctic. Results from our previous investigation of beluga whale mitochondrial DNA (mtDNA) revealed genetic heterogeneity among beluga from different summering locations that was interpreted as representing a high degree of summering site philopatry. However, mtDNA is maternally inherited and does not reflect mating that may occur among beluga from different summering locations in wintering areas or during annual migrations. To test the possibility that breeding occurs among beluga from different summering locations, genetic variability at five nuclear DNA (nDNA) microsatellite loci was examined in the same animals tested in the mtDNA study. Beluga samples ( n = 640) were collected between 1984 and 1994 from 24 sites across North America, mostly during the summer. Whales from the various sites were categorized into eight summering locations as identified by mtDNA analysis, as well as four hypothesized wintering areas: Bering Sea, Hudson Strait (Hudson Strait, Labrador Sea, southwest Davis Strait), Baffin Bay (North Water, east Davis Strait), and St Lawrence River. Microsatellite allele frequencies indicated genetic homogeneity among animals from summering sites believed to winter together but differentiation among whales from some of the wintering areas. In particular, beluga from western North America (Bering Sea) were clearly distinguished from beluga from eastern North America (Hudson Strait, Baffin Bay, and St Lawrence River). Based upon the combined data set, the population of North American beluga whales was divided into two evolutionarily significant units. However, the population may be further subdivided into management units to reflect distinct groups of beluga at summering locations.     

Population structure of North American beluga whales (Delphinapterus leucas) based on nuclear DNA microsatellite variation and contrasted with the population structure revealed by mitochondrial DNA variation

Beluga whales ( Delphinapterus leucas ) in North American waters migrate seasonally between wintering areas in broken pack ice and summering locations in estuaries and other open water areas in the Arctic and sub-Arctic. Results from our previous investigation of beluga whale mitochondrial DNA (mtDNA) revealed genetic heterogeneity among beluga from different summering locations that was interpreted as representing a high degree of summering site philopatry. However, mtDNA is maternally inherited and does not reflect mating that may occur among beluga from different summering locations in wintering areas or during annual migrations. To test the possibility that breeding occurs among beluga from different summering locations, genetic variability at five nuclear DNA (nDNA) microsatellite loci was examined in the same animals tested in the mtDNA study. Beluga samples ( n = 640) were collected between 1984 and 1994 from 24 sites across North America, mostly during the summer. Whales from the various sites were categorized into eight summering locations as identified by mtDNA analysis, as well as four hypothesized wintering areas: Bering Sea, Hudson Strait (Hudson Strait, Labrador Sea, southwest Davis Strait), Baffin Bay (North Water, east Davis Strait), and St Lawrence River. Microsatellite allele frequencies indicated genetic homogeneity among animals from summering sites believed to winter together but differentiation among whales from some of the wintering areas. In particular, beluga from western North America (Bering Sea) were clearly distinguished from beluga from eastern North America (Hudson Strait, Baffin Bay, and St Lawrence River). Based upon the combined data set, the population of North American beluga whales was divided into two evolutionarily significant units. However, the population may be further subdivided into management units to reflect distinct groups of beluga at summering locations.     

Variation in blubber fatty acid composition among marine mammals in the Canadian Arctic

The composition of predator adipose stores can provide important insights into foraging patterns and the ecological relationships among species. We determined the fatty acid (FA) composition of 843 blubber samples from 80 bearded seals (Erignathus barbatus), 33 harbor seals (Phoca vitulina), 239 harp seals (Pagophilus groenlandicus), 32 hooded seals (Cystophora cristata), 281 ringed seals (Phoca hispida), 53 walruses (Odobenus rosmarus rosmarus), 105 beluga whales (Delphinapterus leucas), and 20 narwhals (Monodon monoceros) across the Canadian Arctic to examine patterns of variability among and within species. FA signatures accurately distinguished phocid seals, walruses, and whales. Belugas and narwhals had the most similar FA signatures of any two species, suggesting substantial overlap in their diets, especially in the narwhal‐wintering area off eastern Baffin Island. Among phocid seals, harp and hooded seals had the most similar FA signatures. Bearded seals were most similar to walruses, which was consistent with the benthic feeding habits of both species. Within species, geographic differences in FA signatures were found over both large (>4,000 km) and small (<100 km) spatial scales. Overall, within‐species differences were smaller than among‐species differences. In general, FA signature patterns were consistent with previous studies of the ecology and diets of arctic marine mammals.     

Underwater observations of foraging free-living Atlantic walruses (Odobenus rosmarus rosmarus) and estimates of their food consumption

Food consumption of Atlantic walruses (Odobenus rosmarus rosmarus L.) was quantified by combining underwater observations of feeding with satellite-telemetry data on movement and diving activity. The study was conducted between 31 July and 7 August 2001 in Young Sound (74°N-20°W) in Northeast Greenland. On ten occasions, divers were able to accompany foraging walruses to the sea floor and collect the shells of newly predated bivalves (Mya truncata, Hiatella arctica, Serripes groenlandicus) for determination of number of prey and biomass ingested per dive. Simultaneously, the activity of a 1,200-kg adult male walrus was studied by use of satellite-telemetry during an entire foraging cycle that included 74 h at sea followed by a 23-h rest on land. An average of 53.2 bivalves (SE=5.2, range: 34-89, n=10) were consumed per dive, corresponding to 149.0 g shell-free dry matter (SE=18.9, range: 62.4-253.1 g), or 2,576 kJ per dive (SE=325.2, range: 1,072-4,377 kJ). During the foraging trip, the walrus spent 57% of the time diving to depths of between 6 and 32 m, and it made a total of 412 dives that lasted between 5 and 7 min (i.e. typical foraging dives). If the entire feeding cycle is considered (97 h), the estimated daily gross energy intake was 214 kJ per kg body mass (95% CI: 153-275 kJ), corresponding to the ingestion of 57 kg (95% CI: 41-72 kg) wet weight bivalve biomass per day, or 4.7 (95% CI: 3.3-5.9%) of total walrus body mass. Due to ice cover, walrus access to the plentiful inshore bivalve banks in the area is restricted to the short summer period, where walruses rely on them for replenishing energy stores. It is hypothesised that the documented decrease in the extent and duration of Arctic sea ice may increase food availability for walruses in eastern Greenland in the future.     

Proportion of higher trophic-level prey in the diet of Pacific walruses (Odobenus rosmarus divergens)

During nutritionally stressful situations, Pacific walruses (Odobenus rosmarus divergens) may switch from preying on benthic invertebrates to higher trophic-level prey (HTLP) (e.g., pinnipeds and/or seabirds). We applied a Bayesian mixing model to stable isotope (C and N) data from analyses of various tissues (tongue and lumbar muscle, skin, and liver) to quantify the proportional contribution of HTLP to walruses (n = 293 individuals). The mode contribution of HTLP to walrus diet was ~22 % (±10 %) based on muscle mixing models, which is consistent with results from contaminant studies of Atlantic walruses (Odobenus rosmarus rosmarus), but higher than estimates based on historical stomach content analyses of Pacific walruses. A broader range in the proportion of HTLP (0–60 %) shown by mixing models using stable isotope data from liver and skin of walruses indicated they pursue an opportunistic foraging strategy. Data from the HTLP-consuming walruses were comparable with our stable isotope data of a known “seal-eating” walrus. No significant difference was evident between the estimated contributions of HTLP to the diet of male versus female walruses (P > 0.01). This finding suggests that changes in diet base for walruses are not influenced by the sex of the predator.     

Use of skin biopsies for assessing levels of organochlorines in walruses (Odobenus rosmarus)

Skin and blubber samples of ten adult male Pacific walruses (Odobenus rosmarus divergens) from Alaska were used to investigate the relationship between organochlorine (OC) levels in skin and blubber of individuals. For analyses we selected 11 components that were quantified in the blubber of all individuals: hexachlorocyclohexanes (αHCH and βHCH), the DDT (dichlorodiphenyltrichloroethane) metabolite p,p′DDE, oxychlordane, and 7 individual PCB congeners, 28, 99, 105, 118, 138, 153 and 180. The correlation between the levels in the two types of tissues was significant and the relation was isometric for all components. The regression coefficient between levels in blubber (dependent variable) and levels in skin (independent variable) was different from 1 for only four of the components. The mean levels in the two types of tissues were significantly different for 3 of the 11 chemical components (βHCH, oxychlordane, and PCB28). Although this analysis is based on only ten individuals, we propose that skin samples taken by biopsy darts can be used to monitor OC levels in walruses. In August 1993 skin biopsies were collected from 25 adult male Atlantic walruses (O. r. rosmarus) at haul-out sites in southeastern Svalbard in the Norwegian Arctic and from 28 walruses of different sex and age at haul-out sites at Franz Josef Land in the Russian Arctic. The mean levels of OCs were 2–10 times higher at Svalbard than at Franz Josef Land. The dominant OC component was PCB153 in both areas. A principal component analysis detected differences between areas in OC levels but not in patterns. Since the Franz Josef Land samples were mainly taken from females and young individuals and the Svalbard samples were taken largely from adult males, we believe the differences in tissue OC levels observed from these areas can be explained by differences in sex and age of the walrus sampled. Comparable organochlorine levels in skin samples from walruses from other areas are not available. However, compared to the corresponding OC levels found in walrus blubber in other areas, the OC levels from Svalbard and Franz Josef Land are higher. The high levels of OCs in walruses from Svalbard and Franz Josef Land may be a combined effect of high pollution level in the environment and seal-eating habits. In the present study we show that it is possible to use skin biopsies taken by a non-destructive method to assess OC levels in walruses. Accepted: 24 October 1999     

Mating ecology of beluga (Delphinapterus leucas) and narwhal (Monodon monoceros) as estimated by reproductive tract metrics

Narwhal and beluga whales are important species to Arctic ecosystems, including subsistence hunting by Inuit, and little is understood about their mating ecology. Reproductive tract metrics vary across species in relation to mating strategy, and have been used to infer mating ecology. Reproductive tracts from beluga and narwhal were collected between 1997 and 2008 from five beluga stocks and two narwhal stocks across the Canadian Arctic. Tract length for males and females, relative testes mass for males, and tusk length for male narwhal were measured. We assessed variation relative to species, body size, stock, maturity, and season. Significant variation was found in testes mass across month and stock for beluga, and no significant difference between stock or date of harvest for narwhal. Beluga had significantly larger testes relative to body size than narwhal, suggesting they were more promiscuous than narwhal. A significant relationship was found between narwhal tusk length and testes mass, indicating the tusk may be important in female mate choice. No significant differences were found between narwhal and beluga reproductive tract length for males or females. The mating systems suggested for narwhal and belugas by our results mean the two species may respond differently to climate change.     

Viral and bacterial serology of free-ranging Pacific walrus

Serum or heparinized plasma samples were obtained between 1994 and 1996 from 20 male and 20 female adult free-ranging Pacific walrus (Odobenus rosmarus divergens) from St. Lawrence Island and Round Island, Alaska. Samples were screened for antibodies to some potentially pathogenic bacteria and viruses. No sample had detectable antibody to Brucella spp. Three of 40 (8%) had low antibody titers to Leptospira interrogans serovars. Phocine distemper virus antibodies were not detected. Serologic responses to one or more caliciviruses (San Miguel sea lion virus 12 or vesicular exanthema of swine serotypes E54, F55, G55, 1934B) were detected in 18% (seven of 40) walrus. Antibodies to one or more subtypes of influenza A virus (H10, N2, N3, N5, N6, N7) were detected in 21% (eight of 38). Periodic screening of free-ranging populations for exposure to infectious diseases has become an important component of bio-monitoring programs to facilitate understanding and detecting trends in marine mammal populations.     

Antibodies to marine caliciviruses in the Pacific walrus (Odobenus rosmarus divergens Illiger)

Sera from 155 Pacific walruses (Odobenus rosmarus divergens Illiger), sampled in the Chukchi Sea during the summer of 1983, were tested for serum neutralizing (SN) antibodies to six marine calicivirus serotypes. Serotypes tested included San Miguel sea lion virus (SMSV) types 1, 5, 8, and 10, previously isolated from northern fur seals (Callorhinus ursinus Linné) in the Bering Sea; walrus calicivirus (WCV), previously isolated from walrus feces collected off sea ice in the Chukchi Sea; and Tillamook calicivirus (TCV), a bovine isolate from Oregon of suspected marine origin. No antibodies were found to SMSV-1, SMSV-10, or TCV. Antibodies to SMSV-5 were found in two animals (titers 1:20 and 1:160); antibodies to SMSV-8 were found in four animals (all 1:20); and antibodies to WCV were found in one animal (titer 1:40). Antibodies to WCV have been found in the Pacific walrus previously; however, this represents the first report of antibodies to any of the SMSV serotypes in this marine mammal.     

Inter-annual variability in the proportional contribution of higher trophic levels to the diet of Pacific walruses

Pacific walruses (Odobenus rosmarus divergens) depend on Arctic sea ice as a resting and foraging platform; however, recent years have seen unprecedented seasonal reductions in ice extent. Previous researchers proposed that during unfavorable ice conditions, walruses might prey on other pinnipeds. To examine this hypothesis, we analyzed carbon and nitrogen stable isotope ratios of muscle from walruses (n = 155) sampled from the Bering and Chukchi seas during 2001–2010. We used a Bayesian stable isotope mixing model to examine the proportional contribution of higher trophic level prey (HTLP) (e.g., seals, seabirds) to walrus diets and extrapolated a tissue-specific turnover rate to compare diet of individuals over time. Mode HTLP across years was 19 % ± 8. Results indicate a significant decrease (P < 0.05) in the reliance on HTLP during 2008–2009 (mode HTLP 13 %), one of two sampling periods that experienced great seasonal loss of pan-Arctic sea ice (the other being 2007–2008 with mode HTLP of 23 %). We also reveal intra-annual fluctuations in the contribution of HTLP to the diet of a walrus sampled in 2011 with seal remains in its stomach through high-resolution sectioning along a whisker length. Our findings suggest that walruses forage opportunistically as a result of multiple environmental factors and that sea ice extent alone does not drive consumption of HTLP.     

Results and evaluation of a survey to estimate Pacific walrus population size, 20061

In spring 2006, we conducted a collaborative U.S.–Russia survey to estimate abundance of the Pacific walrus (Odobenus rosmarus divergens). The Bering Sea was partitioned into survey blocks, and a systematic random sample of transects within a subset of the blocks was surveyed with airborne thermal scanners using standard strip‐transect methodology. Counts of walruses in photographed groups were used to model the relation between thermal signatures and the number of walruses in groups, which was used to estimate the number of walruses in groups that were detected by the scanner but not photographed. We also modeled the probability of thermally detecting various‐sized walrus groups to estimate the number of walruses in groups undetected by the scanner. We used data from radio‐tagged walruses to adjust on‐ice estimates to account for walruses in the water during the survey. The estimated area of available habitat averaged 668,000 km² and the area of surveyed blocks was 318,204 km². The number of Pacific walruses within the surveyed area was estimated at 129,000 with 95% confidence limits of 55,000–507,000 individuals. Poor weather conditions precluded surveying in other areas; therefore, this value represents the number of Pacific walruses within about half of potential walrus habitat.