Anthony S. Fischbach

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.

Molecular Method for Determining Sex of Walruses

Abstract We evaluated the ability of a set of published trans-species molecular sexing primers and a set of walrus-specific primers, which we developed, to accurately identify sex of 235 Pacific walruses (Odobenus rosmarus divergens). The trans-species primers were developed for mammals and targeted the X- and Y-gametologs of the zinc finger protein genes (ZFX, ZFY). We extended this method by using these primers to obtain sequence from Pacific and Atlantic walrus (O. r. rosmarus) ZFX and ZFY genes to develop new walrus-specific primers, which yield polymerase chain reaction products of distinct lengths (327 and 288 base pairs from the X- and Y-chromosome, respectively), allowing them to be used for sex determination. Both methods yielded a determination of sex in all but 1–2% of samples with an accuracy of 99.6–100%. Our walrus-specific primers offer the advantage of small fragment size and facile application to automated electrophoresis and visualization.

Spatial genetic structure and asymmetrical gene flow within the Pacific walrus

Abstract Pacific walruses (Odobenus rosmarus divergens) occupying shelf waters of Pacific Arctic seas migrate during spring and summer from 3 breeding areas in the Bering Sea to form sexually segregated nonbreeding aggregations. We assessed genetic relationships among 2 putative breeding populations and 6 nonbreeding aggregations. Analyses of mitochondrial DNA (mtDNA) control region sequence data suggest that males are distinct among breeding populations (ΦST = 0.051), and between the eastern Chukchi and other nonbreeding aggregations (ΦST = 0.336–0.449). Nonbreeding female aggregations were genetically distinct across marker types (microsatellite FST = 0.019; mtDNA ΦST = 0.313), as was eastern Chukchi and all other nonbreeding aggregations (microsatellite FST = 0.019–0.035; mtDNA ΦST = 0.386–0.389). Gene flow estimates are asymmetrical from St. Lawrence Island into the southeastern Bering breeding population for both sexes. Partitioning of haplotype frequencies among breeding populations suggests that individuals exhibit some degree of philopatry, although weak. High levels of genetic differentiation among eastern Chukchi and all other nonbreeding aggregations, but considerably lower genetic differentiation between breeding populations, suggest that at least 1 genetically distinct breeding population remained unsampled. Limited genetic structure at microsatellite loci between assayed breeding areas can emerge from several processes, including male-mediated gene flow, or population admixture following a decrease in census size (i.e., due to commercial harvest during 1880–1950s) and subsequent recovery. Nevertheless, high levels of genetic diversity in the Pacific walrus, which withstood prolonged decreases in census numbers with little impact on neutral genetic diversity, may reflect resiliency in the face of past environmental challenges.

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

ABSTRACT 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. Summary: Age-specific muscle biochemistry of the major locomotor muscles in walruses enables estimates to be made of their aerobic dive limits and theoretical bottom times relative to age and dive depth.

Walrus haul-out and in water activity levels relative to sea ice availability in the Chukchi Sea

An animals energetic costs are dependent on the amount of time it allocates to various behavioral activities. For Arctic pinnipeds, the time allocated to active and resting behaviors could change with future reductions in sea ice cover and longer periods of open water. The Pacific walrus (Odobenus rosmarus divergens) is a large Arctic pinniped that rests on sea ice or land between foraging trips to feed on the seafloor. We used behavioral data collected from radiotagged walruses in the Chukchi Sea (2008–2014) in a Bayesian generalized linear mixed effects model to estimate the probability a walrus was in water foraging, in water not foraging, or hauled out, as a function of environmental covariates. The probability of a walrus being in water increased with wind speed and decreased with air temperature, and the probability a walrus was foraging, given it was in water, increased with available benthic macrofaunal biomass. The probability of each behavior was also related to the nature and availability of haul-out substrates. The amount of time walruses spent in water foraging and hauled out was greatest when only sea ice was available, which typically occurs when walruses occupy feeding areas during summer and early autumn. This situation may be most energy efficient for walruses because it allows the highest proportion of in water energy expenditure to be allocated to foraging. Conversely, the amount of time walruses spent in water foraging and hauled out was lowest when only land was available, which typically occurs in late autumn, in years when walruses were constrained to land haul-outs because sea ice was absent over the continental shelf.

Evaluation of a method using survey counts and tag data to estimate the number of Pacific walruses (Odobenus rosmarus divergens) using a coastal haulout in northwestern Alaska

Increased periods of sparse sea ice over the continental shelf of the Chukchi Sea in late summer have reduced offshore haulout habitat for Pacific walruses (Odobenus rosmarus divergens) and increased opportunities for human activities in the region. Knowing how many walruses could be affected by human activities would be useful to conservation decisions. Currently, there are no adequate estimates of walrus abundance in the northeastern Chukchi Sea during summer–early autumn. Estimating abundance in autumn might be possible from coastal surveys of hauled out walruses during periods when offshore sea ice is unavailable to walruses. We evaluated methods to estimate the size of the walrus population that was using a haulout on the coast of northwestern Alaska in autumn by using aerial photography to count the number of hauled out walruses (herd size) and data from 37 tagged walruses to estimate availability (proportion of population hauled out). We used two methods to estimate availability, direct proportions of hauled out tagged walruses and smoothed proportions using local polynomial regression. Point estimates of herd size (4200–38,000 walruses) and total population size (76,000–287,000 walruses) ranged widely among days and between the two methods of estimating availability. Estimates of population size were influenced most by variation in estimates of availability. Coastal surveys might be improved most by counting walruses when the greatest numbers are hauled out, thereby reducing the influence of availability on population size estimates. The chance of collecting data during peak haulout periods would be improved by conducting multiple surveys.