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.     

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.     

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).     

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     

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     

AERIAL CENSUS OF PACIFIC WALRUSES IN THE CHUKCHI SEA, 1985

The U.S. Fish and Wildlife Service conducted a survey of the walruses in the pack ice of the Chukchi Sea between 16 September and 2 October 1985, as part of a joint effort with the Soviet Union to estimate the size of the Pacific walrus population. American observers conducted censuses from two aircraft along randomly selected north–south lines over the pack ice. The observers counted walruses within a constant viewing angle that corresponded to a total strip width of 1.38 km at an altitude of 152 m.
In nine days of flying, 15,312 walruses were observed, of which 10,140 were on 5,764 km² of census strips. Few walruses were observed east of 161°W longitude or west of 170°W longitude, hence the census effort was stratified. Walrus concentrations between 161° and 170° shifted slightly westward during the 2-wk duration of the censuses. The differences among days in observed walrus density were due to changes in the numbers of walruses on the ice within 37 km of the ice edge. The number of observable walruses in pack ice of the eastern Chukchi Sea was estimated to be 62,177 (SD = 19,480), based on censuses conducted on 29 and 30 September and 1 October. At that time there were also at least 15,238 in Bristol Bay, Bering Sea. The Soviets counted 39,572 on the shores of the western Chukchi and Bering seas and estimated 115,531 in pack ice of the western Chukchi Sea. Summing U.S. and Soviet estimates, the total population of Pacific walruses in 1985 was 232,518. This number was comparable with earlier estimates from censuses conducted jointly by the U.S. and the Soviets. However, information on fraction hauled out, segregation, and movements is needed for more precise estimates.     

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.     

Estimation of walrus populations on sea ice with infrared imagery and aerial photography

Population sizes of ice‐associated pinnipeds have often been estimated with visual or photographic aerial surveys, but these methods require relatively slow speeds and low altitudes, limiting the area they can cover. Recent developments in infrared imagery and its integration with digital photography could allow substantially larger areas to be surveyed and more accurate enumeration of individuals, thereby solving major problems with previous survey methods. We conducted a trial survey in April 2003 to estimate the number of Pacific walruses (Odobenus rosmarus divergens) hauled out on sea ice around St. Lawrence Island, Alaska. The survey used high altitude infrared imagery to detect groups of walruses on strip transects. Low altitude digital photography was used to determine the number of walruses in a sample of detected groups and calibrate the infrared imagery for estimating the total number of walruses. We propose a survey design incorporating this approach with satellite radio telemetry to estimate the proportion of the population in the water and additional low‐level flights to estimate the proportion of the hauled‐out population in groups too small to be detected in the infrared imagery. We believe that this approach offers the potential for obtaining reliable population estimates for walruses and other ice‐associated pinnipeds.     

Regional walrus abundance estimate in the United States Chukchi Sea in autumn

Human activities (e.g., shipping, tourism, oil, gas development) have increased in the Chukchi Sea because of declining sea ice. The declining sea ice itself and these activities may affect Pacific walrus (Odobenus rosmarus divergens) abundance; however, previous walrus abundance estimates have been notably imprecise. When sea ice is absent from the eastern Chukchi Sea, walruses in waters of the United States usually rest together onshore at a single Alaska coastal haulout, where they can be surveyed more easily than when they rest on dispersed offshore ice floes. We estimated the number of walruses on land (herd size) at this haulout from 13 unoccupied aircraft system (UAS) surveys flown within a 10-day period in each of 2018 and 2019. We estimated population size of walruses using the haulout over the course of the surveys by combining herd size data with data from satellite-linked transmitters that indicated whether tagged walruses were in or out of water during each survey. Our estimates of the population size of walruses using the haulout during each year's survey period were similar to each other and more precise than historical walrus abundance estimates: posterior means (95% credibility intervals) were 166,000 (133,000–201,000) for 2018 and 189,000 (135,000–251,000) for 2019. Auxiliary observations support using these estimates to represent the size of the population using the eastern Chukchi Sea in autumn during the surveyed years. Our study site was the only substantial Chukchi Sea coastal haulout in the United States during the survey periods and study-specific tracking data (consistent with known distribution and movement patterns) indicated tagged walruses remained in eastern Chukchi waters during the survey periods. In addition, the imagery, telemetry, and analytical methods developed for this study advance the prospect for precise range-wide walrus population size estimates.     

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.     

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.     

COMPARISON OF METHODS USED TO ESTIMATE NUMBERS OF WALRUSES ON SEA ICE

The US and former USSR conducted joint surveys of Pacific walruses on sea ice and at land haul-outs in 1975, 1980, 1985, and 1990. One of the difficulties in interpreting results of these surveys has been that, except for the 1990 survey, the Americans and Soviets used different methods for estimating population size from their respective portions of the sea ice data. We used data exchanged between Sovier and American scientists to compare and evaluate the two estimation procedures and to derive a set of alternative estimates from the 1975, 1980, and 1985 surveys based on a single consistent procedure. Estimation method had only a small effect on total population estimates because most walruses were found at land haul-outs. However, the Sovievr method is subject to bias that depends on the distribution of the population on the sea ice and this has important implications for interpreting the ice portions of previously reported surveys for walruses and other pinniped species. We recommend that the American method be used in future surveys. Future research on survey methods for walruses should focus on other potential sources of bias and variation.     

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.     

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.     

Phylogeographic origin and genetic diversity of dunlin <Emphasis Type="Italic">Calidris alpina</Emphasis> in Svalbard

We investigated the genetic structure of the presumably small (10–100 pairs) and isolated dunlin (Calidris alpina) population breeding in Svalbard, and compared it with similar data recently published from several dunlin populations in the western Palearctic and East Greenland. Using mitochondrial and nuclear DNA markers, as well as data on bill lengths, we sought to infer the phylogeographic origin of Svalbard dunlins and assess their within-population level of genetic diversity. Only dunlins with haplotypes of the European mtDNA clade (EUR) were found in Svalbard, indicating a close resemblance to dunlin populations in East Greenland and Iceland. Microsatellite data for Svalbard dunlins, as well as their short bills, also supported a western origin. The Svalbard population did not show signs of inbreeding or reduced levels of genetic diversity compared to other investigated populations, which suggests that the population was recently founded or is currently subject to considerable gene flow.     

Identifying a reliable blubber measurement site to assess body condition in a marine mammal with topographically variable blubber,the Pacific walrus

Pacific walruses may be unable to meet caloric requirements in the changing Arctic ecosystem, which could affect body condition and have population‐level consequences. Body condition has historically been monitored by measuring blubber thickness over the xiphoid process (sternum). This may be an unreliable condition index because blubber at other sites along the body may be preferentially targeted to balance energetic demands. Animals in aquaria provided an opportunity for controlled study of how blubber topography is altered by caloric intake. Morphology, body mass, blubber thickness (21 sites), and caloric intake of five mature, nonpregnant, nonlactating female walruses were measured monthly (12 month minimum). Body condition (mass × standard length^?1) was described by a model that included caloric intake and a seasonal effect, and scaled positively with estimates of total blubber mass. Blubber thicknesses (1.91–10.69 cm) varied topographically and were similar to values reported for free‐ranging female walruses. Body condition was most closely related to blubber thickness measured dorsomedially in the region of the anterior insertion of the pectoral flippers (shoulders); sternum blubber thickness was a relatively poor indicator of condition. This study demonstrates the importance of validating condition metrics before using them to monitor free‐ranging populations.     

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.