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.     

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.     

Geographic variation of PCB congeners in polar bears (Ursus maritimus) from Svalbard east to the Chukchi Sea

We present data on geographic variation in polychlorinated biphenyl (PCB) congeners in adult female polar bears (Ursus maritimus) from Svalbard eastward to the Chukchi Sea. Blood samples from 90 free-living polar bears were collected in 1987-1995. Six PCB congeners, penta to octa chlorinated (PCB-99, -118, -153, -156, -180, -194), were selected for this study. Differences between areas were found in PCB levels and congener patterns. Bears from Franz Josef Land (11,194 ng/g lipid weight) and the Kara Sea (9,412 ng/g lw) had similar DPCB levels and were higher than all other populations (Svalbard 5,043 ng/g lw, East Siberian Sea 3,564 ng/g lw, Chukchi Sea 2,465 ng/g lw). Svalbard PCB levels were higher than those from the Chukchi Sea. Our results, combined with earlier findings, indicate that polar bears from Franz Josef Land and the Kara Sea have the highest PCB levels in the Arctic. Decreasing trends were seen eastwards and westwards from this region. Of the congeners investigated in the present study, the lower chlorinated PCBs are increasing and the high chlorinated PCBs are decreasing from Svalbard eastward to the Chukchi Sea. Different pollution sources, compound transport patterns and regional prey differences could explain the variation in PCB congener levels and patterns between regions.     

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.     

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     

Benthic scavengers collected by baited traps in the high Arctic

Baited traps were deployed in the coastal areas of Svalbard (0–175 m), Franz Josef Land (15 and 20 m), Northeast Greenland (6 and 20 m) and Ellesmere Island (2–145 m). The samples were dominated by Lysianassidae (Amphipoda). In the most extensively sampled locality (Hornsund, Svalbard), differences between the necrophagic fauna of the outer and inner basins were observed. Clear depth separations of closely related species of the Anonyx and Onisimus genera were found between 0 and 175 m in Hornsund. Anonyx nugax and Onisimus edwardsi were the only species found in all of the examined localities. Accepted: 14 January 2000     

RESPONSE OF PACIFIC WALRUSES TO DISTURBANCES FROM CAPTURE AND HANDLING ACTIVITIES AT A HAUL-OUT IN BRISTOL BAY, ALASKA

Observations were made on herds of the Pacific walrus ( Odobenus rosmarus divergens ) to study their response during the capturing and handling of adult males in summer 1995 at a haul-out at Cape Peirce in southwestern Alaska. Three behaviors (alertness, displacement, and dispersal) were quantified from 16 capture sessions. Herd sizes ranged from 622 to 5,289 walruses. Handling of an immobilized walrus consisted of attempts to attach telemetry devices to the tusks and collect various biological samples. Handling activities resulted in an average of about 10-fold or greater levels of behavior in alertness, displacement, and dispersal than during precapture and darting periods. High levels of behavior usually occurred within the first 45 min of handling. In 8 of 10 capture sessions, walruses returned to predisturbance levels of behavior within 40 min of cessation of the handling disturbance. Alertness and displacement were moderately and negatively correlated with herd size during the handling period, which may reflect an effect of a threshold distance from the point of disturbance to responding individuals. Observations of walruses tagged with VHF radio transmitters indicated that the activities from a given capture session did not preclude tagged walruses from using the haul-out over a subsequent 11 -wk monitoring period. Moreover, non-tagged walruses continued to extensively use the haul-out during and after the period in which capture sessions were conducted.     

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.     

The number of walruses (Odobenus rosmarus) in Svalbard in summer

Walruses were on the verge of extinction in Svalbard prior to becoming totally protected in 1952. The present study attempts to determine the number of walruses in Svalbard in summer based on aerial, ground and boat surveys. In 1993, 741 walruses were observed, almost all of which were males. This population, which is shared with the neighbouring Russian archipelago Franz Joseph Land, is believed to consist of an equal number of females. We therefore conclude that the shared population consists of a minimum of about 1450 walruses older than 2 years and an unknown number of calves.     

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.     

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.     

Geographical variation in the breeding biology of the Purple Sandpiper Calidris maritima

Studies in Iceland (66°N) and Svalbard (78°N), combined with the results of previous work, allowed geographical comparisons of different aspects of the breeding biology of the Purple Sandpiper Calidris maritima , testing predictions that they may be related to latitude and altitude (here taken as surrogates for climatic severity). The breeding density was lowest in the polar deserts of Franz Josef Land (82°N) and highest in coastal Arctic Tern Sterna paradisaea colonies in Iceland where the terns perhaps afforded protection from predators. There was no difference in the percentages of 1-year-old (first-year) birds in the breeding populations in Svalbard and Iceland. Time of breeding was related to both latitude and altitude, being later at higher latitudes and altitudes. In Iceland, larger males and males over 1 year old nested earlier than small males and 1-year-old males, respectively. Relaying occurred in Iceland but it was not observed in Svalbard. There were no geographical differences in clutch size. Egg size was related to female size in Svalbard. There were also geographical differences in egg size associated with female size (wing length), with the largest eggs in Iceland and smallest in south Norway. Males usually attended broods and the rates at which females deserted their broods were similar in Iceland and Svalbard. Chick growth rates were similar for Iceland and Franz Josef Land. Thus, in breeding biology, Purple Sandpipers varied geographically only in breeding density, time of nesting and egg size, which was associated with female size.     

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.     

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.     

Polar bear (<Emphasis Type="Italic">Ursus maritimus</Emphasis>) maternity den distribution in Svalbard,Norway

The aim of the current study was to identify maternity denning areas and to describe aspects of denning behaviour in polar bears (Ursus maritimus) in Svalbard, Norway. Den observations made during aerial and ground surveys (n = 553), observations of recently emerged family groups made during capture and tagging of bears (n = 206), and dens recorded by satellite telemetry (n = 76 dens) were used to describe the distribution of dens in the Barents Sea region. Data were collected during March, April, and May from 1972 to 2010, and while most dens were abandoned during the first 2 weeks of April, there was substantial variation. The highest number of dens was recorded on the islands in the eastern and northern parts of Svalbard with fewer dens found further west on the island Spitsbergen. Females with cubs-of-the-year or their tracks indicated some denning in southern, eastern, and northern parts of Spitsbergen. Telemetry also revealed that females captured in Svalbard and the Barents Sea den in Franz Josef Land, Russia. The majority of dens (62%) in Svalbard were located on land within ca. 1 km of the shore with the most inland den 19 km from shore. Heavy exploitation of polar bears before 1973, when a hunting moratorium was implemented, likely led to reduced denning in some areas. Our observations of den distribution indicate that denning is now more widespread in the archipelago than 50 years ago and reflects a reestablishment of denning areas following years of protection.     

Identification of motivational state in adult male Atlantic walruses inferred from changes in movement and diving behavior

Motivational changes in animals are likely to be detectable retrospectively through observed changes in behavior. Breeding represents one of the strongest motivational states in mammals, and its timing is often tied to a seasonally optimal suite of environmental and physical conditions. While seasonal changes in behavior may be directly observable in some species, for others that breed cryptically or in difficult to access areas, detecting behavioral changes may only be feasible using data collected remotely. Herein, we explore whether behavioral changes can be used to infer motivational state for a wild, free‐ and wide‐ranging high arctic marine mammal, adult male Atlantic walruses (Odobenus rosmarus rosmarus). Using satellite‐relayed location and dive data from 23 adult male walruses instrumented in the Svalbard Archipelago, we identify seasonal movement to discrete geographic regions deep into winter pack ice. Adult male walrus diving behavior underwent marked seasonal movements between geographical areas that coincided with changes in light regime. At offshore wintering sites adult males (n = 4) shifted from a summer pattern of deep, long benthic dives to much shallower diving. Some males performed similar shallow, winter dive behavior at coastal locations (n = 12) suggesting that breeding might also occur around the coast of Svalbard. However, interpretation of behavioral changes of these coastal individuals was challenging. The presumed breeding sites at the winter off‐shore locations were situated in areas where polynyas are known to occur, making them a predictable resource even if they are located deep inside the winter pack‐ice. We demonstrate that remotely collected behavioral data can be used to identify seasonally explicit changes in the behavior of cryptic species.     

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.     

Records of the gadoid fish <Emphasis Type="Italic">Arctogadus glacialis</Emphasis> (Peters, 1874) in the European Arctic

The Arctogadus glacialis is endemic to the Arctic Ocean and its apparently disjunct circumpolar distribution range from the Siberian coast through the Chukchi Sea and the Canadian Arctic to the shelf off NE Greenland. Records of A. glacialis are scarce in the European Arctic and here we present all available and reliable records of the species in the area. Altogether, 296 specimens of A. glacialis are reported from 53 positions in the European Arctic during the period 1976–2008. The specimens were registered off Iceland and the Jan Mayen Island, northwest and northeast of Svalbard, northeast in the Barents Sea, and south and east off Franz Josef Land. The additional records show that A. glacialis display a circumpolar and more continuous distribution than described before. In the European Arctic, A. glacialis has been caught at 155–741 m depth with the highest abundance at 300–400 m. We therefore suggest that A. glacialis is more associated to the continental shelves surrounding the Arctic Ocean than previously thought. The length–weight relation of A. glacialis is similar across the European Arctic.     

On a typical Jurassic sauropterygian from the Upper Triassic of Wilczek Land (Franz Josef Land,Arctic Russia)

A new elasmosaurid genus and species, Alexeyisaurus karnoushenkoi, from the Lower-Middle Norian (Wilczek Formation) of the Upper Triassic of the island of Wilczek Land (Franz Josef Land) is described based on an incomplete skeleton. The new form combines characters of typical late plesiosaurs and structural features unique to this form. It is probably the earliest representative of typical late sauropterygians (plesiosaurs).     

Morphological and thermal properties of mammalian insulation: the evolutionary transition to blubber in pinnipeds

Carnivora includes three independent evolutionary transitions to the marine environment: pinnipeds (seals, sea lions, and walruses), sea otters, and polar bears. Among these, only the pinnipeds have retained two forms of insulation, an external fur layer and an internal blubber layer for keeping warm in water. In this study we investigated key factors associated with the transition to the use of blubber, by comparing blubber characteristics among the pinnipeds. Characteristics included gross morphology (blubber thickness), fat composition (fatty acid profiles, percentage lipid, and water), and thermal conductivity. Sea lions, phocids, and walrus, which have lower fur densities than fur seals, have thicker blubber layers than fur seals (P < 0.001). Comparisons of lipid content, water content, and fatty acid composition indicated significant differences in the composition of the inner and outer regions of the blubber between groups (P < 0.001), consistent with the hypothesis that phocids and sea lions utilize the outer layer of their blubber primarily for thermal insulation, and the inner layer for energy storage. Fur seals, by contrast, rely more on their fur for thermal insulation, and utilize their blubber layer primarily for energy storage. Comparing across carnivore species, differences in total insulation (fur and/or blubber) are influenced substantially by body size and habitat, and to a lesser extent by latitudinal climate. Overall, these results indicate consistent evolutionary trends in the transition to blubber and evidence for convergent evolution of thermal traits across lineages. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.