Dentition in Pacific walrus (Odobenus rosmarus divergens) calves of the year

The oral cavity and teeth were examined in Pacific walrus calves of the year that died or were harvested on Cape Vancarem and Koluchin Island, the Chukchi Sea, in September to October 2010. The order of gingival eruption of teeth, the resultant dental formula in calves of the year and 1.5-year-old animals, and the dental formula after alveolar eruption of teeth were determined. The number of teeth after gingival eruption in calves of the year did not depend on their body length and sex. The loss of some teeth, mainly incisors, was recorded in individuals aged 5?C6 months. Already at early developmental stages, functional teeth were found to differ significantly from other (rudimentary) teeth in their weight, total size, and the size of apical foramen. Some individuals had milk teeth, which were either gingivally erupted or partly remained within soft tissues. Cases of aberration in dentition that require additional studies were recorded.     

Demography of the Pacific walrus (Odobenus rosmarus divergens) in a changing Arctic

The Pacific walrus (Odobenus rosmarus divergens) is a candidate to be listed as an endangered species under United States law, in part, because of climate change‐related concerns. While the population was known to be declining in the 1980s and 1990s, its recent status has not been determined. We developed Bayesian models of walrus population dynamics to assess the population by synthesizing information on population sizes, age structures, reproductive rates, and harvests for 1974–2015. Candidate models allowed for temporal variation in some or all vital rates, as well as density dependence or density independence in reproduction and calf survival. All selected models indicated that the population underwent a multidecade decline, which began moderating in the 1990s, and that annual reproductive rate and natural calf survival rates rose over time in a density‐dependent manner. However, selected models were equivocal regarding whether the natural juvenile survival rate was constant or decreasing over time. Depending on whether juvenile survival decreased after 1998, the population growth rate either increased during 1999–2015 or stabilized at a lesser level of decline than seen in the 1980s. The probability that the population was still declining in 2015 ranged from 45% to 87%.     

Projected status of the Pacific walrus (Odobenus rosmarus divergens) in the twenty-first century

Extensive and rapid losses of sea ice in the Arctic have raised conservation concerns for the Pacific walrus (Odobenus rosmarus divergens), a large pinniped inhabiting arctic and subarctic continental shelf waters of the Chukchi and Bering seas. We developed a Bayesian network model to integrate potential effects of changing environmental conditions and anthropogenic stressors on the future status of the Pacific walrus population at four periods through the twenty-first century. The model framework allowed for inclusion of various sources and levels of knowledge, and representation of structural and parameter uncertainties. Walrus outcome probabilities through the century reflected a clear trend of worsening conditions for the subspecies. From the current observation period to the end of century, the greatest change in walrus outcome probabilities was a progressive decrease in the outcome state of robust and a concomitant increase in the outcome state of vulnerable. The probabilities of rare and extirpated states each progressively increased but remained <10% through the end of the century. The summed probabilities of vulnerable, rare, and extirpated (P(v,r,e)) increased from a current level of 10% in 2004 to 22% by 2050 and 40% by 2095. The degree of uncertainty in walrus outcomes increased monotonically over future periods. In the model, sea ice habitat (particularly for summer/fall) and harvest levels had the greatest influence on future population outcomes. Other potential stressors had much smaller influences on walrus outcomes, mostly because of uncertainty in their future states and our current poor understanding of their mechanistic influence on walrus abundance.     

Pacific walrus (Odobenus rosmarus divergens):Differential prey digestion and diet

Stomach content data from 798 Pacific walruses ( Odobenus rosmarus divergens ) collected during 1952–1991 were analyzed using a method that evaluates the stage of digestion of prey remains. Non-molluscan prey taxa were not well represented in previous interpretations of walrus diet due to digestion biases. Stomach contents least affected by digestion (fresh stomachs) contained more prey taxa than stomachs of an unknown or more digested state. Bivalves, gastropods, and polychaete worms were the most frequent prey items in both the Bering and Chukchi seas, although bivalves occurred more frequently in stomachs from the Bering Sea and gastropods occurred more frequently in stomachs from the Chukchi Sea. Male and female walruses consumed essentially the same prey when in the same location. Using only fresh stomachs collected between 1975 and 1985, there was no significant difference between the proportion that contained mostly bivalves and the proportion that contained non-bivalve prey items. Earlier interpretations of a change in walrus diet in this period compared to the prior two decades may have been due to digestion as well as sampling biases. Current climatic changes may affect walrus's access to diverse, productive shallow water feeding areas.     

Demography of the Pacific walrus (Odobenus rosmarus divergens): 1974–2006

Global climate change may fundamentally alter population dynamics of many species for which baseline population parameter estimates are imprecise or lacking. Historically, the Pacific walrus is thought to have been limited by harvest, but it may become limited by global warming‐induced reductions in sea ice. Loss of sea ice, on which walruses rest between foraging bouts, may reduce access to food, thus lowering vital rates. Rigorous walrus survival rate estimates do not exist, and other population parameter estimates are out of date or have well‐documented bias and imprecision. To provide useful population parameter estimates we developed a Bayesian, hidden process demographic model of walrus population dynamics from 1974 through 2006 that combined annual age‐specific harvest estimates with five population size estimates, six standing age structure estimates, and two reproductive rate estimates. Median density independent natural survival was high for juveniles (0.97) and adults (0.99), and annual density dependent vital rates rose from 0.06 to 0.11 for reproduction, 0.31 to 0.59 for survival of neonatal calves, and 0.39 to 0.85 for survival of older calves, concomitant with a population decline. This integrated population model provides a baseline for estimating changing population dynamics resulting from changing harvests or sea ice.     

Trend in Pacific walrus (Odobenus rosmarus divergens) tusk asymmetry, 1990–2014

We used the basal circumference of Pacific walrus (Odobenus rosmarus divergens) tusks (upper canine teeth, n = 21,068 pairs) to estimate fluctuating asymmetry (FA1 index) from 1990 to 2014. The mean difference in circumference between paired tusks was ?0.006 (SEM = 0.002) cm and approximately normally distributed. Measurement error was 0.6 (0.02)%, similar between biologists and lay persons (P = 0.83), and ≤15% of FA1. Tusk FA1 was greatest in 1990 then declined by 56% (P = 0.0001) through 2014. Male and female trends differed (P = 0.0001) and male FA1 was 40% greater (P = 0.0001) and the rate of decline 28% steeper (P = 0.3) than females. A quartic polynomial model (r² = 0.66, w_i = 0.685) fit the trend for female data better than simpler forms, whereas a linear model (r² = 0.55, w_i = 0.693) was a better fit for male data. Walrus tusk FA1 reflected periods when the population was stressed due to food limitations and then recovered, and perhaps when females began to experience the loss of preferred sea ice habitat in summer and FA1 is an easily monitored indicator. More work is needed to confirm the link between FA1, individual fitness, and adaptive potential.     

Faecal DNA amplification in Pacific walruses (Odobenus rosmarus divergens)

Dietary information is critical for assessing the population status of seals, sea lions and walruses—and is determined for most species of pinnipeds using non-invasive methods. However, diets of walruses continue to be described from the stomach contents of dead individuals. Our goal was to assess whether DNA could be extracted from the faeces of Pacific walruses (Odobenus rosmarus divergens) collected at haulout sites, and whether potential prey species or taxa could be amplified from that DNA. We extracted DNA from 70 faecal samples collected from ice pans in the Bering Sea during the spring of 2008 and 2009 (with between 4.6 and 308.9 ng/μl of DNA in every sample). We also extracted DNA from 12 potential prey species or taxa collected by bottom-grabs in 2009 to identify positive controls for primers and to test the ability of previously published taxon-specific and species-specific primers to correctly identify the prey using conventional PCR. We tested primers that successfully amplified DNA from the tissue of at least one potential prey species or taxon on all 70 walrus faecal samples. We found that two sets of primers successfully amplified many of the potential prey species or taxa using DNA from their tissue, and that one of these primer sets produced positive amplification in 4 of the 70 faecal samples. The band size that was produced for prey organisms and in the faecal samples was consistent with expectations, although prey identities were not verified with sequencing. Our pilot study demonstrates that DNA can be successfully extracted and amplified from walrus faeces, providing a stepping stone towards describing the diets of walruses from faecal DNA.     

Persistent toxic substances in the muscles and liver of the pacific walrus Odobenus rosmarus divergens Illiger, 1815 from the Bering Sea

This study reports data on the contents of organochlorine pesticides and heavy metals in the muscles and livers of eight individuals (five males and three females) of the Pacific walrus Odobenus rosmarus divergens Illiger, 1815, which were captured in the summer of 2011 in Mechigmensky Bay of the Bering Sea. Pesticides, namely α-, β-, and γ-isomers of hexachlorocyclohexane (HCH), dichlorodiphenyltrichloroethane (DDT) and its metabolites dichlorodiphenyldichloroethane (DDD) and dichlorodiphenyldichloroethylene (DDE), as well as the heavy metals Cd, Pb, and Hg were found in all samples studied. The total concentration of organochlorine pesticides in the muscles varied from 200 to 5700 ng/g lipid weight and in the liver from 4900 to 90300 ng/g lipid weight. The concentrations of cadmium, lead, and mercury were 0.04–6.7, 0.13–0.76, and 0.03–0.40 μg/g dry weight, respectively. On the whole, the contents of organochlorine pesticides and heavy metals in the organs of the Pacific walrus were lower compared to those in marine mammals from other regions of the World Ocean.     

The method of age determination of the Pacific walrus (Odobenus rosmarus divergens) using the layered structure of functional teeth

The functional teeth of Pacific walruses that died or were harvested in the Retkyn Spit, Enmelen village, Kolyuchin Island, Vankarem Cape, Enurmino village, and Chegitun River region in 2005, 2007–2008, and 2010–2011 were examined. The definite structure was investigated in animals of their first year of life. The presence of a milk layer in the tooth dentine can be considered as a mark that all the cement layers have been preserved, and the age determination of an individual is precise. The annual increment of dentine significantly changed with age in different parts of the tooth (buccal, lingual, and central), and the annual growth of dentine decreased every year. The growth rate of the upper jaw teeth was significantly higher, and the duration of their growth was much longer than that in the lower jaw teeth. The wearing of dentine and cement layers was unequal in different parts of tooth. Several recommendations for choosing a tooth for the determination of the walrus’s age and for the estimation of age using the layered tooth structure are given.     

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.     

Models of tongue movement in the walrus (Odobenus rosmarus)

Three hypothetical models of tongue movement of the walrus during suction feeding are examined. These models encompass the entire range of simple tongue retraction movements possible by examining 1) movement of the tongue directly to the rear following the curvature of the palate, 2) to the rear and ventrally in a straight line, and 3) ventrally in a straight line. The percent of muscular force available from the hyoglossus, genioglossus, and styloglossus that could be applied toward retraction as predicted by each model is calculated. The resistance that the tongue would provide during retraction is calculated using projected tongue areas and is combined with the above data from the muscles to provide an estimate of the percent of the total available force that is needed to retract the tongue for each model. A separate examination of the direction of tongue-induced wear striations on the palatal and lingual aspects of the teeth is used to help support or reject the three models. The model where the tongue is moved directly to the rear is supported by studies of both muscle force and tooth wear. In the mammalian groups that were compared to the walrus, there is a great deal of interspecific variation in movements of the tongue during suction feeding; no two groups can be considered to have identical stereotyped tongue movements.     

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.     

The influence of ice conditions on terrestrial haulouts of the pacific walrus Odobenus rosmarus divergens Illiger, 1815 in the Gulf of Anadyr,Bering Sea

The boundaries of the area of appropriate depths for feeding by the Pacific walrus were determined. Based on the ice maps for the period of 1997–2011, the long-term dynamics of ice breakup and formation in the Gulf of Anadyr (Bering Sea) were analyzed. There was a tendency towards a reduction in the ice-cover period in the Gulf of Anadyr, especially during 2007–2011. In 2008, the number and duration of stay of walruses at terrestrial haulout sites on Meeskyn Spit Island and on Retkyn Spit continued to decrease. Walruses also used other haulouts located in the Gulf of Anadyr at Cape Gek and Cape Retkyn. The age and sex structure of the walrus herd changed: the proportions of adult males and young animals decreased, while the proportion of females with calves increased.     

Carnivora: the primary structure of the Pacific Walrus (Odobenus rosmarus divergens, Pinnipedia) hemoglobin

The primary structure of the alpha- and beta-chains of the hemoglobin from the Pacific Walrus (Odobenus rosmarus divergens, Pinnipedia) is presented. Sequence analysis revealed only one hemoglobin component whereas two bands were found in polyacrylamide gel electrophoresis. The globin chains were separated by high-performance liquid chromatography and the sequences determined by automatic liquid- and gas-phase sequencing of the chains and their tryptic peptides. The alpha-chains show 20 and the beta-chains 12 exchanges compared to the corresponding human chains. In the alpha-chains one heme- and two alpha 1/beta 1-contacts were exchanged whereas in the beta-chains one alpha 1/beta 1-, one alpha 1/beta 2-and one heme-contact are substituted. Compared to Harbour Seal (Phoca vitulina) the Walrus hemoglobin shows 9 amino-acid replacements in the alpha-chains and 5 in the beta-chains. The relation between Pinnipedia and Arctoidea is discussed.     

A review of the genetic relationships of Atlantic walrus (Odobenus rosmarus rosmarus) east and west of Greenland

Studies of the genetic variation involving allozymes, mitochondrial and nuclear DNA (microsatellites) in walruses (Odobenus rosmarus) were reviewed. In addition, the genetic relationships of a total of 211 Atlantic walruses, O. r. rosmarus, from 5 sampling areas west and east of Greenland were studied using 12 nuclear DNA-microsatellite loci and restriction fragment length polymorphism obtained from the ND1, ND2 and ND3/4 segments of the mitochondrial DNA (mtDNA). At the mtDNA level, no divergence was observed among the three sampling areas east of Greenland (i.e. East Greenland, Svalbard and Franz Josef Land), whereas areas west of Greenland (i.e. Northwest and West Greenland) showed some differentiation. The genetic variation at the microsatellite loci grouped the individuals into four sub-populations: Northwest Greenland, West Greenland, East Greenland and a common Svalbard-Franz Josef Land sub-population. A significant correlation between genetic distance and geographic distance between the sampling areas (isolation-by-distance effect) was detected, especially at the mtDNA level. At a small-scale phylo-geographical level, the mtDNA data indicated that Atlantic walruses have diverged into two major groups: one northwest (i.e. in the North Water) and one east of Greenland (i.e. an East Greenland-Svalbard-Franz Josef Land group), whereas the haplotype distribution in the West Greenland sample reflected a mixture of both these groups. The microsatellite data supported a general grouping of walruses to the west and east of Greenland.     

Spatial variation and low diversity in the major histocompatibility complex in walrus (Odobenus rosmarus)

Increased global temperature and associated changes to Arctic habitats will likely result in the northward advance of species, including an influx of pathogens novel to the Arctic. How species respond to these immunological challenges will depend in part on the adaptive potential of their immune response system. We compared levels of genetic diversity at a gene associated with adaptive immune response [Class II major histocompatibility complex (MHC), DQB exon 2] between populations of walrus (Odobenus rosmarus), a sea ice-dependent Arctic species. Walrus was represented by only five MHC DQB alleles, with frequency differences observed between Pacific and Atlantic populations. MHC DQB alleles appear to be under balancing selection, and most (80 %; n = 4/5) of the alleles were observed in walruses from both oceans, suggesting broad scale differences in the frequency of exposure and diversity of pathogens may be influencing levels of heterozygosity at DQB in walruses. Limited genetic diversity at MHC, however, suggests that walrus may have a reduced capacity to respond to novel immunological challenges associated with shifts in ecological communities and environmental stressors predicted for changing climates. This is particularly pertinent for walrus, since reductions in summer sea ice may facilitate both northward expansion of marine species and associated pathogens from more temperate regions, and exchange of marine mammals and associated pathogens through the recently opened Northwest Passage between the Atlantic and Pacific Oceans in the Canadian high Arctic.     

The influence of feeding,enrichment, and seasonal context on the behavior of Pacific Walruses (Odobenus rosmarus divergens)

Though some research exists concerning general behavior and activity patterns of Walruses in zoos or aquariums, less is known about how these patterns change in response to various environmental and temporal contexts. This study presents two studies assessing behavioral changes in relation to feeding period, object enrichment (OE), and season in a social group of four Pacific Walruses at the New York Aquarium. Study 1 examined behavior in relation to feeding context (nonfeed, prefeed, postfeed); data were collected over a three‐week period, resulting in 47 observation sessions for each feeding context. Study 2 examined behavior in relation to OE and season; data were collected in two phases resulting in 12 enrichment and 9 no‐enrichment (NE) observation sessions (Phase 1), and 21 enrichment and 18 NE observation sessions (Phase 2). Study 1 showed that after feeding, oral behavior increased while social behavior and total swim frequency decreased. In Study 2, both swim frequency and social behavior were found to interact with OE and phase, while oral behavior remained constant across all conditions. As in the wild, both studies found all animals to be swimming the majority of the time. Though every animal spent much of its swim time engaged in an Individual Swimming Pattern (ISP), both studies showed that the proportion of ISP (in relation to total time swimming) remained stable across all contexts, suggesting a potential functional role of the ISPs. These results are discussed in light of the ongoing debate over the role of stereotypies in welfare assessment. Zoo Biol 29:397–404, 2010. © 2009 Wiley‐Liss, Inc.