Body size of male Atlantic walruses (Odobenus rosmarus rosmarus) from Svalbard

Morphometric data are given for 41 adult male Atlantic walruses ( Odobenus rosmarus rosmarus) immobilized at Svalbard. Total body weight ranged from 960 to 1450 kg. Standard body length ranged from 258 to 380 cm, while curvilinear tusk length ranged from 17 to 65 cm. Immobilized animals were selected for their large size. The sample is therefore biased towards larger animals. Based on regression equations developed for walruses caught in Greenland, the weights of Svalbard walruses were estimated from standard body length and axillary girth. Estimated weights ranged from 632 to 1883 kg. The present data suggest that Atlantic walruses at Svalbard can attain the same body size as Pacific ( O. r. divergens ) and Laptev walruses (O. r. laptevi).     

Diving and haul-out patterns of walruses Odobenus rosmarus on Svalbard

Nine male walruses were equipped with dive recording devices in Svalbard to investigate walrus diving and haul-out behaviour in late summer. Dive information on 6,018 dives was collected by 3 satellite linked dive recorders. Additional dive information on 7,769 dives was obtained from 3 time depth recorders. The deepest dive recorded was 67 m, but mean depth of foraging dives was 22.5 m. The longest-lasting dive recorded was 24 min, but mean duration of foraging dives was 6 min. The walruses, on average, spent 56 h in the water followed by 20 h hauled out on land.     

Body growth in Atlantic walruses (Odobenus rosmarus rosmarus) from Greenland

Morphometric data were collected from 105 Atlantic walruses ( Odobenus rosmarus rosmarus ) in northwestern Greenland in the periods 1977–78 and 1989–91. Of these 21 walruses were subjected to a detailed study on body composition.
The asymptotic maximum standard body length of Atlantic walruses in NW Greenland was 269 cm for females and 314 cm for males. This is similar to Pacific walruses, but significantly longer than Atlantic walruses from Hudson Bay. Despite this, walruses from NW Greenland apparently do not attain the same total body mass as Pacific walruses ( O.r.divergens ).
The asymptotic maximum body weights for walruses in NW Greenland were estimated to be 720 kg for females and 1114 kg for males.
Total body surface area was proportional to the two-thirds power of total body weight.
The percentage proportion of blubber and viscera were both negatively correlated to body mass, while skin and muscles constituted a nearly fixed proportion. On average, blubber constituted 19% of total body mass of adult females, 15% of adult males and 24% of subadults of both genders. The average walrus consisted of 18% blubber, 12% skin, 12% viscera and 58% blood, muscle and skeleton. Muscles were estimated to constitute 44% of total body weight.
Allometric functions for weight of internal organs relative to body mass are presented.     

Levels of cadmium and mercury in the hair of Atlantic walruses (Odobenus rosmarus rosmarus) from Svalbard,Norway

Hair samples of 15 adult male Atlantic walruses (Odobenus rosmarus rosmarus) collected from anaesthetized individuals at Svalbard, Norway, were analysed for cadmium and total mercury. The mean level of cadmium was 0.860 ± 0.321 μg/g dry weight (median = 0.811, range = 0.349–1.51 μg/g dry weight) and the mean level of mercury was 0.235 ± 0.100 μg/g dry weight (median = 0.251, range = 0.121–0.424 μg/g dry weight). Levels of cadmium and mercury in hair of walruses from other areas are not known. Both cadmium and mercury levels in hair of walruses from Svalbard are relatively low compared to the levels found in the hair of other marine mammal species. It has been documented from a number of marine species, including marine mammals such as ringed seals and polar bears, that both cadmium and mercury levels at Svalbard are lower than in other areas. It is uncertain as to what degree levels in hair reflect levels in internal organs in walruses. In rare and highly endangered species or populations tissue samples can be difficult to collect. In walruses, it is possible to collect hair from anaesthetized individuals or at the haul-out sites during moult, to monitor heavy metal levels of the population. Accepted: 6 December 1998     

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.     

Resting and swimming metabolic rates in juvenile walruses (Odobenus rosmarus)

Changes in Arctic ice conditions have raised concerns regarding potential impacts on energy expenditure and food requirements of walruses. Modeling the repercussions of environmental changes requires accurate species-specific measures of bioenergetic expenditures. This is particularly true for walruses, who have a unique anatomy and foraging ecology from other pinnipeds. This study measured resting metabolic rate (RMR) and subsurface swimming metabolism in two juvenile walruses over a 13-month period. The walruses had relatively low RMR compared to studies of other young pinnipeds. RMR was greater for the male than the female, as expected given its larger size; the reverse was true on a mass-specific basis. There was also considerable variability in RMR for each walrus during the year that could not be accounted for by changes in body mass. Metabolism while swimming was about twice RMR, and locomotor costs were higher than generally predicted for other marine mammals. The lower calculated swimming efficiency may reflect the fact that walruses are not “high velocity” pursuit predators. The estimates of metabolic expenditure obtained in this study for young walruses are invaluable for quantifying the energetic consequences of behavioral changes induced by environmental shifts in the wild.     

Haul-out and diving activity of male Atlantic walruses (Odobenus rosmarus rosmarus) in NE Greenland

During August-September 1989 and 1990, movements, haul out and dive activity of male Atlantic walruses ( Odobenus rosmarus rosmarus L.) were studied at a terrestrial haul-out site situated in an inshore foraging area in NE Greenland at 76± 30' N. Data were collected from direct observations of a group of about 50 males during August, including walruses that could be individually identified from natural markings, and from tracking of 8 adults equipped with satellite-linked radio transmitters during August-September. In both years, instrumented walruses hauled out for a total of 29.3% of the sampling time. In 1989, when ice floes were available for hauling out, the walruses spent 11% of the time on ice, whereas in 1990, when ice was absent from the study area, they only hauled out on land. Duration of haul-out periods, which did not differ between months or years, averaged 11 h (0.46 d) on ice (S.D. = 5.9, range: 1–29 h, n = 19 periods), and 38 h (1.6 d) on land (S.D. = 11.7, range: 13–64 h, n = 43). The walruses mainly hauled out during the afternoon and evening. Numbers hauling out on land during August were negatively correlated with wind direction, precipitation (rain) and wind-chill. In 1989, the duration of periods of absence from the terrestrial haul-out site (i.e. presumed foraging trips) averaged 206 h or 8.5 d (S.D. = 106.9, range: 48–412 h, n = 13), whereas, in 1990, such trips averaged only 81 h or 3.4 d (S.D. = 37.9, range: 24-156 h, n = 24), reflecting that walruses used the haul-out site more frequently when ice was absent. Direct observations of foraging walruses showed that they were submerged about 81% of the time.     

Aerial survey of Atlantic walruses (Odobenus rosmarus rosmarus) in the Pechora Sea, August 2011

Basic knowledge of the population biology of Atlantic walruses throughout the eastern parts of their range, including the Pechora Sea, is scarce or nonexistent. Herein, we present the first estimate of walrus numbers from the Pechora Sea based on an aerial survey of 2,563?km of coastline, using a combination of infrared techniques and digital imagery. Hauled out walruses were found at three sites (Vaygach Island and two sites on Matveyev Island). A total of 968 animals were counted on aerial photographs; all of the animals appeared to be males. Crude measurements of dorsal curvilinear lengths of a subset of the photographed animals (N?=?504) showed that many were adults, but 14.5?% belonged to younger age classes (shorter than 225?cm). Using an adjustment factor developed for male walruses in Svalbard, to account for animals at sea during the survey, the number of walruses occupying this area was estimated to be 3,943 (95?% CI, 3,605–4,325). No females with calves were seen in this survey, implying that the population that uses the Pechora Sea during summer has a distributional area that is larger than the survey area. Extensive oil exploration, development and production are currently taking place in the Pechora Sea. Risks posed to walruses and their prey by these industrial activities should be assessed immediately, and the genetic delineations of this population should be clarified.     

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.     

Temporal association of serum progesterone concentrations and vaginal cytology in walruses (Odobenus rosmarus)

Concentrations of serum estradiol-17β and progesterone were monitored in six female walruses using an enzyme immunoassay. Progesterone concentrations increased from March to May in females aged 6 y or older, and subsequently declined (October). No significant elevation of estradiol-17β concentration was detected before an elevation of progesterone concentration. Vaginal smears from four females were examined with Papanicolaou staining. In all females, most epithelial cells were basophilic intermediate-superficial cells; no color change from basophilic to eosinophilic of the cells was detected. Meanwhile, the percentage of anucleate cells in vaginal smears reached its highest value before the elevation of progesterone concentration, followed by an increase in the percentage of leukocytes. We inferred that the change in populations of anucleate cells and leukocytes in vaginal smears reflected ovarian status and CL formation in female walruses.     

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     

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.     

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.     

Killer whales (Orcinus orca) hunting for walruses (Odobenus rosmarus divergens) near Retkyn Spit, Chukotka

Group hunting by killer whales for walruses was observed in August 18, 2008, in the littoral area (3 km from the haulout of walruses, Retkyn Spit, Chukotka). The group of killer whales consisted of seven adults (one adult male did not participate in attacks) and two calves. Based on prey type, these killer whales were mammal-eating. The total duration of their hunt activity was not less than 95 min. The hunt consisted of three phases. The first phase was an attack on the group of walruses and choice of individual prey; the second phase was attacks on the chosen walrus; and the third (final) phase was a decrease in activity of killer whales and leaving group with walrus from sea shore. The main behavioral patterns of killer whales during the hunt were discerned. Two killer whales tried to kill walruses by chasing them and jumping out of the water on the shore. The video analysis of the ??attack phase?? showed that killer whales made 55 attacks on the walrus during 17.3 min. On average, each killer whale attacked the walrus seven times. The attack tactics of killer whales, the number of movements, and the location of killer whales (adults and calves) relative to each other and to the walrus were described. Well coordination of their movements and group actions was observed.     

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.     

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.