Walrus Research Articles

The mussel path – Using the contaminant tracer, Ecotracer, in Ecopath to model the spread of pollutants in an Arctic marine food web

As the polar ice cap is receding, shipping in the Arctic seas becomes easier, and both destination and Atlantic–Pacific transit shipping is expected to increase. Thereby, the risk of accidents increase. Immediate negative impacts are expected from oil spills through the acute mortality for marine organisms, especially from heavy fuel oil (HFO). Marine Diesel oil (MDO) is therefore suggested as a preferable fuel for ships operating in Arctic waters. However, Polycyclic Aromatic Hydrocarbons (PAHs) are toxic components in both types of fuel, are highly bioavailable and can transfer up the food chain. A spill of MDO following a shipwreck could therefore have impacts beyond the spill site and long after the diesel has spread and evaporated. We model the spread of PAHs from a fictitious spill of MDO in the Pechora Sea (South Eastern Barents Sea) using the contaminant tracer module Ecotracer, in the Ecopath modelling software. We address the effects on the food-web including long term effects by combining toxicology and food-web modelling. Ecotracer assumes that pollutants follow the biomass passively through the system, and degradation of pollutants is following user specified rates. By combining in natura measurements of PAHs in seawater and in blue mussels (Mytilus edulis) recorded at an accidental MDO spill site, with experiments conducted on the red king crab (Paralithodes camtschaticus) and blue mussels, we derived values as inputs into the model. The Ecotracer predicted that the pollution in the mussels will spread throughout the food-web, especially to the top predators of mussels, king eider (Somateria spectabilis) and Atlantic walrus (Odobenus rosmarus rosmarus) and also from snow crab (Chionoecetes opilio) to seals and toothed whales.

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

Physiological constraints dictate animals' ability to exploit habitats. For marine mammals, it is important to quantify physiological limits that influence diving and their ability to alter foraging behaviors. We characterized age-specific dive limits of walruses by measuring anaerobic (acid-buffering capacity) and aerobic (myoglobin content) capacities of the muscles that power hind (longissimus dorsi) and fore (supraspinatus) flipper propulsion. Mean buffering capacities were similar across muscles and age classes (a fetus, five neonatal calves, a 3month old and 20 adults), ranging from 41.31 to 54.14slykes and 42.00 to 46.93slykes in the longissimus and supraspinatus, respectively. Mean myoglobin in the fetus and neonatal calves fell within a narrow range (longissimus: 0.92-1.68 g 100 g(-1) wet muscle mass; supraspinatus: 0.88-1.64 g 100 g(-1) wet muscle mass). By 3 months post-partum, myoglobin in the longissimus increased by 79%, but levels in the supraspinatus remained unaltered. From 3months post-partum to adulthood, myoglobin increased by an additional 26% in the longissimus and increased by 126% in the supraspinatus; myoglobin remained greater in the longissimus compared with the supraspinatus. Walruses are unique among marine mammals because they are born with a mature muscle acid-buffering capacity and attain mature myoglobin content early in life. Despite rapid physiological development, small body size limits the diving capacity of immature walruses and extreme sexual dimorphism reduces the diving capacity of adult females compared with adult males. Thus, free-ranging immature walruses likely exhibit the shortest foraging dives while adult males are capable of the longest foraging dives.

Thermal reference points as an index for monitoring body temperature in marine mammals.

Background Monitoring body temperature is essential in veterinary care as minor variations may indicate dysfunction. Rectal temperature is widely used as a proxy for body temperature, but measuring it requires special equipment, training or restraining, and it potentially stresses animals. Infrared thermography is an alternative that reduces handling stress, is safer for technicians and works well for untrained animals. This study analysed thermal reference points in five marine mammal species: bottlenose dolphin (Tursiops truncatus); beluga whale (Delphinapterus leucas); Patagonian sea lion (Otaria flavescens); harbour seal (Phoca vitulina); and Pacific walrus (Odobenus rosmarus divergens).ResultsThe thermogram analysis revealed that the internal blowhole mucosa temperature is the most reliable indicator of body temperature in cetaceans. The temperatures taken during voluntary breathing with a camera held perpendicularly were practically identical to the rectal temperature in bottlenose dolphins and were only 1 °C lower than the rectal temperature in beluga whales. In pinnipeds, eye temperature appears the best parameter for temperature control. In these animals, the average times required for temperatures to stabilise after hauling out, and the average steady-state temperature values, differed according to species: Patagonian sea lions, 10 min, 31.13 °C; harbour seals, 10 min, 32.27 °C; Pacific walruses, 5 min, 29.93 °C.ConclusionsThe best thermographic and most stable reference points for monitoring body temperature in marine mammals are open blowhole in cetaceans and eyes in pinnipeds.

The first steps towards generating induced pluripotent stem cells from cryopreserved skin biopsies of marine mammals

A new approach for restoring populations of endangered marine mammal species is to use induced pluripotent stem cells (iPSCs). These cells can be obtained by genetic reprogramming of somatic animal cells, such as skin fibroblasts. From there, the iPSCs can be differentiated into all cell types of the organism, including spermatozoa and oocytes. The skin biopsies of three species of marine mammals, belonging to different orders (the walrus Odobenus rosmarus, the Steller sea lion Eumetopias jubatus and the Irrawaddy dolphin Orcaella brevirostris), were taken and cryopreserved. The fibroblast cultures were obtained from thawed skin biopsies of sea lion and walrus using previously developed methods for human cells. In contrast, Irrawaddy dolphin skin cells did not survive this cell isolation procedure. We had to modify the isolation method to obtain a fibroblast culture from this animal, using mechanical disaggregation only and a higher concentration of serum and antibiotics at the first steps of cultivation. The functionally active fibroblasts will be used in the next step, generating iPSCs. Testing the quality of the fibroblast cultures, we found that the abnormal fibroblasts differed in size and the distribution of filamentous actin and tubulin, compared to normal cells, and demonstrated clear aberrations in the nuclei.

A New Late Miocene Odobenid (Mammalia: Carnivora) from Hokkaido, Japan Suggests Rapid Diversification of Basal Miocene Odobenids.

The modern walrus, Odobenus rosmarus, is specialized and only extant member of the family Odobenidae. They were much more diversified in the past, and at least 16 genera and 20 species of fossil walruses have been known. Although their diversity increased in the late Miocene and Pliocene (around 8–2 Million years ago), older records are poorly known. A new genus and species of archaic odobenid, Archaeodobenus akamatsui, gen. et sp. nov. from the late Miocene (ca. 10.0–9.5 Ma) top of the Ichibangawa Formation, Hokkaido, northern Japan, suggests rapid diversification of basal Miocene walruses. Archaeodobenus akamatsui is the contemporaneous Pseudotaria muramotoi from the same formation, but they are distinguishable from each other in size and shape of the occipital condyle, foramen magnum and mastoid process of the cranium, and other postcranial features. Based on our phylogenetic analysis, A. akamatsui might have split from P. muramotoi at the late Miocene in the western North Pacific. This rapid diversification of the archaic odobenids occurred with a combination of marine regression and transgression, which provided geological isolation among the common ancestors of extinct odobenids.

Uridine Diphosphate-Glucuronosyltransferase (UGT) Xenobiotic Metabolizing Activity and Genetic Evolution in Pinniped Species.

There are various interspecies differences in xenobiotic-metabolizing enzymes. It is known that cats show slow glucuronidation of drugs such as acetaminophen and strong side effects due to the UGT1A6 pseudogene. Recently, the UGT1A6 pseudogene was found in the Northern elephant seal and Otariidae was suggested to be UGT1A6-deficient. From the results of measurements of uridine diphosphate-glucuronosyltransferase (UGT) activity using liver microsomes, the Steller sea lion, Northern fur seal, and Caspian seal showed UGT activity toward 1-hydroxypyrene and acetaminophen as low as in cats, which was significantly lower than in rat and dog. Furthermore, UGT1A6 pseudogenes were found in Steller sea lion and Northern fur seal, and all Otariidae species were suggested to have the UGT1A6 pseudogene. The UGT1 family genes appear to have undergone birth-and-death evolution based on a phylogenetic and synteny analysis of the UGT1 family in mammals including Carnivora. UGT1A2-1A5 and UGT1A7-1A10 are paralogous genes to UGT1A1 and UGTA6, respectively, and their numbers were lower in cat, ferret and Pacific walrus than in human, rat, and dog. Felidae and Pinnipedia, which are less exposed to natural xenobiotics such as plant-derived toxins due to their carnivorous diet, have experienced fewer gene duplications of xenobiotic-metabolizing UGT genes, and even possess UGT1A6 pseudogenes. Artificial environmental pollutants and drugs conjugated by UGT are increasing dramatically, and their elimination to the environment can be of great consequence to cat and Pinnipedia species, whose low xenobiotic glucuronidation capacity makes them highly sensitive to these compounds.

Bioaccumulation of persistent organochlorine pesticides (OCPs) by gray whale and Pacific walrus from the western part of the Bering Sea

The feeding habits of a gray whale (Eschrichtius robustus) and a Pacific walrus (Odobenus rosmarus divergens), caught from the western Bering Sea in the summers of 2010 and 2011, have been studied, and concentration of persistent organochlorine pesticides (OCPs) in their organs determined. The total OCP concentration (∑HCH+∑DDT) in muscles and liver of the gray whales varies from 297 to 3581 and from 769 to 13,808ng/g lipids, respectively. The total OCP concentration (∑HCH+∑DDT) in muscles and liver of the Pacific walruses varies from 197 to 5659 and from 4856 to 90,263ng/g lipids, respectively. The specifics of diet as a source of pesticide accumulation in these two marine mammal species are discussed.

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

AbstractMotivational 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.

Comparison of managed care and wild walrus source characteristics

Male Pacific walruses perform acoustic displays while in rut. The purpose of these displays is unknown but are hypothesized to be for territory defense or mate advertisement. Understanding source characteristics will allow the estimation of perceptibility by conspecifics. The displays occur in the Bering Sea in late winter where direct human observation is difficult. Working with an animal in managed care provided the ability to make direct observations of a male producing breeding vocalizations and the direct calculation of source level. Source characteristics from recordings of managed care and wild walruses were analyzed. The mean peak source level of the impulsive knocks produced by the managed care male was 183 dB (re: 1 μPa). The mean peak source level from the wild recordings was 177 dB (re: 1 μPa). For both wild and managed care vocalizations, a significant relationship between ambient noise level and source level was identified. An increase of approximately 5 dB in source level was found for an i...

Demographic inferences using short-read genomic data in an approximate Bayesian computation framework: in silico evaluation of power, biases and proof of concept in Atlantic walrus.

Approximate Bayesian computation (ABC) is a powerful tool for model-based inference of demographic histories from large genetic data sets. For most organisms, its implementation has been hampered by the lack of sufficient genetic data. Genotyping-by-sequencing (GBS) provides cheap genome-scale data to fill this gap, but its potential has not fully been exploited. Here, we explored power, precision and biases of a coalescent-based ABC approach where GBS data were modelled with either a population mutation parameter (θ) or a fixed site (FS) approach, allowing single or several segregating sites per locus. With simulated data ranging from 500 to 50000 loci, a variety of demographic models could be reliably inferred across a range of timescales and migration scenarios. Posterior estimates were informative with 1000 loci for migration and split time in simple population divergence models. In more complex models, posterior distributions were wide and almost reverted to the uninformative prior even with 50000 loci. ABC parameter estimates, however, were generally more accurate than an alternative composite-likelihood method. Bottleneck scenarios proved particularly difficult, and only recent bottlenecks without recovery could be reliably detected and dated. Notably, minor-allele-frequency filters - usual practice for GBS data - negatively affected nearly all estimates. With this in mind, we used a combination of FS and θ approaches on empirical GBS data generated from the Atlantic walrus (Odobenus rosmarus rosmarus), collectively providing support for a population split before the last glacial maximum followed by asymmetrical migration and a high Arctic bottleneck. Overall, this study evaluates the potential and limitations of GBS data in an ABC-coalescence framework and proposes a best-practice approach.

Abundance and distribution of pacific walrus <i>Odobenus rosmarus divergens</i> in vicinity of Cape Serdtse-Kamen in 2009-2013

Results of the first special research of pacific walrus Odobenus rosmarus divergens at Cape Serdtse-Kamen conducted in the 2009-2011 and 2013 are presented. The coastal haulout place looks as series of relatively isolated areas (up to 36) located alongshore about 20 km from Cape Serdtse-Kamen to the Ikalyurynveem River mouth. Initially the walruses were on the northwestern edge of the haulout place between Cape Serdtse-Kamen and Cape Sulyn but expanded southeastward with increasing of their number. In the years with early and intense ice melting, the walruses come to the area in early September, and the haulout forms to the end of the first 10-day period of this month. In conditions of late ice retreat, as in the 2013, the walruses on the whole come to Cape Serdthse-Kamen later. The maximum number of walruses in this area was observed in 2009 (about 98,000 animals) and 2010 (118,500). This is the record number of walruses ever recorded simultaneously on one coastal haulout in the species range. Almost the whole Pacific population gathers in the vicinity of Cape Serdtse-Kamen for feeding in late autumn and continues to feed there until freezing of the Chukchi Sea. In spite of obvious significance of the area at Cape Serdtse-Kamen for the pacific walrus population, there still aren’t any special measures of their protection in this part of the coast when they are ashore - the measures are definitely necessary to prevent intensive economic activities in this key for the walruses area and to save the coastal haulout for traditional use of native people.

Satellite traceology: experience of using in the interests of mammalogy for protection of ice-associated marine mammals

Advanced technology of satellite traceology is presented on the base of using the synthetic aperture radar (SAR) with high space and deep resolution installed for the first time onboard Soviet space station Almaz and now functioning successfully onboard the satellites Envisat and RADARSAT. The SAR traceology is the central issue of the satellite eco-criminology and concerns to theory of the traces formation, variability, and preservation in undamaged state. Ships coordinates, their traces and complexity of ice routing are fixed by the method of satellite charting, and their possible influence on ice-associated marine mammals is assessed. Numerous examples of SAR satellite control on navigation in stormy weather and ships’ presence in ice with different compactness, origin and age are overviewed, as well as impact of travel facilities on ice-associated animals welfare. Some incidents potentially dangerous for marine mammals are considered for the Gulf of Finland (Baltic Sea). Parameters of the ice cover, as openings, large fractures, and polynyas made by icebreakers in this area were determined on the base of the satellite SAR data, comprehensive digital ice maps with scheme of real ships’ routes in the ice were prepared, and the routes correspondence with ESIMO demands was assessed. The satellite SAR survey in the Kandalaksha Bay (White Sea) allowed to fix that winter navigation in the White Sea provoked formation of fractures and polynyas and reduced the sea ice area suitable for reproduction of greenland seals. For validation of this situation, airborne control was organized aboard the aircraft L-410 «Nord» that provided panoramic and IR images of ship channel through the rookeries of the seals in time of their reproduction. Besides, the aircraft made observations of pacific walruses behavior in the Bering Sea which were superposed with the satellite SAR and passive microwave survey in frame of the project «Pacific Walrus». Traceological control of the sea ice in the Anadyr Bay (Bering Sea) allowed to determine the size of openings in the ice and to assess the influence of tide and wind on welfare of ice-associated marine mammals. Some cases of infringement the regulation of navigation were revealed by means of the satellite SAR traceology.

On factors for separation of the stock of beluga whale <i>Delphinapterus leucas</i> in the Anadyr Gulf of the Bering Sea

New eco-ethological data on the Anadyr stock of beluga whales are presented based on 14-year (2000-2013) studies by methods of satellite tracking telemetry, multi-spectral aerial surveys and genetic analysis, as well as traditional and ecosystem-based approaches. Possible factors of the separate stock formation in the Anadyr Gulf are discussed, as geographic isolation, environmental conditions, foraging resources, elimination of the beluga whales, and biological isolation (on the results of genetic analysis). The beluga whales are well-adapted to oceanographic conditions and ice regime of the Arctic waters that is realized in their relationship with edge of the sea ice, ability to live in both salt and fresh waters (they enter rivers), and versatility in feeding. Their ice-associating, high sociality, and white protective coloration provide protection against killer whales and other predators and their eury-halinity allows to expand the feeding grounds both to the sea and rivers and hunt for both freshwater, anadromous and marine prey. The food spectrum of beluga whales in the Anadyr estuary is very wide and includes 12 fish species and 1 crustacean species. In the summer-autumn season, the salmons are the most significant part of the diet, as chum salmon Oncorhynchus keta , pink salmon O. gorbuscha and arctic char Salvelinus malma , whereas arctic smelt Osmerus mordax dentex , saffron cod Eleginus gracilis , sculpins Cottidae sp., anadyr whitefish Coregonus anaulorum , and siberian whitefish Coregonus sardinella are presented in the diet to a lesser extent. There is supposed on the base of episodic net surveys, that the beluga whales feeding in marine areas is more diverse, and the most important their prey in the sea are walleye pollock Theragra chalcogramma , pacific cod Gadus macrocephalus , flounders Pleuronectidae sp., sculpins Cottidae sp., capelin Mallotus villosus catervarius , polar cod Boreogadus saida , halibuts Pleuronectidae sp., stingrays Bathyraja , herring Clupeidae sp., and several species of decapod crustaceans Hyppolitidae sp. and Lithodidae sp. Natural enemies of beluga whales are killer whale Orcinus orca , polar bear Ursus maritimus , and predatory form of pacific walrus Odobenus rosmarus . Rather large number of beluga whales are eliminated by the sea ice, up to 80 animals annually, that exceeds in several times the aboriginal landings in Chukotka. Parasitic fauna of the beluga whale is still unclear and requires special investigations; the cases of epizooty are unknown in the Anadyr Gulf. Philopatry is proper to the Anadyr beluga whales, with preference of the Anadyr estuary and the Anadyr Gulf as their habitats, that is explained by optimal for them environments and high food capacity. Unique adaptation of beluga whales to the environments of the Anadyr estuary is their ultrasonic vocalization in the high-turbidity waters. Genetic analysis shows a differentiated distribution of the beluga whales haplotypes in different areas of the North Pacific, so the stock of the Anadyr Gulf is significantly isolated in the post-glacial times. Low anthropogenic pressure on beluga whales in the Anadyr Gulf allows them to maintain their stock there.

Prolonged chemical restraint of walrus (<i>Odobenus rosmarus</i>) with etorphine supplemented with medetomidine

Physiological studies involving the use of isotopic water required chemical restraint of free-ranging walruses (Odobenus rosmarus) for several hours. In August 2000, six male walrus (total body mass: 1050–1550 kg) were immobilized in East Greenland by remote delivery of 8.0–9.8 mg of etorphine and subsequently restrained for up to 6.75 h by administration of medetomidine. The effects of etorphine were reversed with 10–24 mg diprenorphine. After termination of the etorphine-induced apnoea, lasting an average of 15.8 min (SD = 9.7, range = 9.5–35.2 min, n = 6), the animals were initially given 10–20 mg medetomidine intramuscularly. The initial dose was further augmented by 5 mg at intervals of 5 min. In two cases, when medetomidine was administered through a catheter inserted in the extradural vein, the animal became instantly apnoeic and regained respiratory function only after intravenous injection of the prescribed dose of the antagonist atipamezole and of the respiratory stimulant doxapram. After an average of 3.5 hours of immobilisation, rectal temperature began to increase and it is conceivable that this is the factor that will ultimately limit the duration of immobilisation. The animals became conscious and fully mobile shortly after an intravenous injection of a dose of atipamezole approximately twice the mass of the total dose of medetomidine given during the procedure followed by 400 mg of doxapram. It is concluded that medetomidine appears to be a suitable drug for chemical restraint of walruses for time-consuming procedures following initial immobilisation by etorphine. With animals of total body mass around 1,000–1,500 kg, the drug should be given intramuscularly in 10–20 mg increments (total mass 10–60 mg) until the breathing rate falls to approximately 1 min-1. At this level, breathing is maintained and animals do not respond to touch or injection.

Behavior and body mass changes of a mother and calf Pacific walrus (Odobenus rosmarus divergens) during the suckling period.

The behavior of a 13-year-old female Pacific walrus and her first calf is described during the first 7 months of the 19-month suckling period. The calf vocalized before 62% of suckling sessions. The mother immediately responded after 84% of vocalizations; after 44% she allowed a suckling session. The number of suckling sessions per 24-h period decreased from on average nine in the second week after birth to two by the thirteenth week. Thereafter, the number of suckling sessions fluctuated between 3 and 5/day. The average suckling session duration increased from 4 min/day to around 17 min/day. The average effective suckling time per session increased from on average 2 to 10 min. The number of breaks decreased during the study period from around 40 to 20 per session. The mother spent on average 47% of her time resting, the calf 44%. The remaining time was spent in locomotion, and suckling, playing, investigating, and looking. Mother and calf spent on average 51% of time indoors. The mother spent 43% of her time in the water, the calf 39%. The calf's body mass at birth was 55 kg; it increased to 178 kg at 27 weeks. The mother did not eat for the first 5 days after delivery, and she ate less than usual during the next 6 days. The mother's mass eventually stabilized at 1024 kg (>before gestation). The calf was not given formula and was weaned onto fish. This is the first detailed description of a captive Pacific walrus's suckling period. It could benefit the husbandry of future captive-born walruses, and may elucidate the behavior of wild walruses during suckling.

Abundance of walruses in Eastern Baffin Bay and Davis Strait

Walruses (Odobenus rosmarus) are exploited for subsistence purposes in West Greenland. However, current information about the abundance of walruses subject to harvest in eastern Baffin Bay subject to harvest has been unavailable despite being critical for maintaining sustainable catch levels. Three visual aerial surveys were conducted in 2006 (21 March to 19 April 2006), 2008 (3 to 12 April) and 2012 (24 March to 14 April) to estimate the number of walruses on the wintering grounds in eastern Baffin Bay and Davis Strait. Data on the fraction of walruses that were submerged below a 2m detection threshold during the surveys were obtained from 24 walruses instrumented with satellite-linked-time-depth-recorders in northern Baffin Bay in May-June 2010-2012. An availability correction factor was estimated at 36.5% (cv=0.08) after filtering of data for an observed drift of the pressure transducer of more than 2.5 m. The surveys resulted in walrus abundance estimates that were corrected for walruses submerged below a detection threshold and for walruses that were missed by the observers. The estimates of abundance were 1,105 (cv=0.31, 95% CI 610-2,002) in 2006, 1,137 (0.48, 468-2,758) in 2008 and 1,408 (0.22, 922-2,150) in 2012.

Movements of walruses (<i>Odobenus rosmarus</i>) between Central West Greenland and Southeast Baffin Island, 2005-2008

Between 2005 and 2008, 31 walruses were tagged with Argos satellite transmitters at their wintering grounds at Store Hellefiske Banke, Central West Greenland (N=23), and at their summering grounds off the coast of Southeast Baffin Island, Canada (N=8). Two male walruses moved along the Greenland coast from Store Hellefiske Banke north to Disko Banke, where contact was lost. Two other males went further north, up to the Upernavik area. Contact was lost with one of these tags, but the other animal travelled southward again and went towards SE Baffin Island. Eight of the tagged walruses moved between West Greenland and Baffin Island, demonstrating a connection between walruses at these sites. Walruses left the Store Hellefiske Banke during the period 7 April to 17 May and on average used 7 days to swim the 400 km across Davis Strait. The migration routes were quite similar; they travelled over the shallowest areas at the narrowest part of the strait. The timing of the spring dispersal and migration towards Canada was closely linked to the extent and timing of the retreat of the pack ice edge. One flipper tagged male that was marked off South Baffin Island was recovered in a hunt on Store Hellefiske Banke, documenting that the reverse migration also occurs. Off West Greenland satellite tagged walruses spent a lot of time around the Store Hellefiske Banke (55.0o-56.5o W), using this shallow area as feeding grounds irrespective of the ice cover in this area. Partial sexual segregation was observed. Despite a tendency in West Greenland for males to occur farther offshore, farther from the ice edge and at greater depths, only their preference for denser ice cover (64% ice cover) differed significantly (P=0.019) from the habitat preferences of females (52% ice cover). Coast dispersal was more condensed and the segregation between males and females was more pronounced during autumn along the Southeast Baffin Island. Females remained farther north (P<0.001) and farther east (P=0.006), and males were more often located offshore in areas with greater water depths (P=0.024). Males had also had larger home range than females during both seasons.

Management of Atlantic walrus (<i>Odobenus rosmarus rosmarus</i>) in the arctic Atlantic

We review the management of Atlantic walruses (Odobenus rosmarus rosmarus) past and present in the four range states—Canada, Greenland, Norway and Russia—which have permanent populations of Atlantic walruses. Populations in all four countries have been depleted, although the extent of depletion is not well known. Inuit in Arctic Canada and Greenland hunt Atlantic walruses for subsistence while they have been protected at Svalbard (Norway) since 1952 and in the western Russian Arctic since 1956. Since the second half of the 20th Century Canada and Greenland have increased protection of their walrus. Generally the number of walruses landed in Canada is governed by the number of hunters and/or people in the settlement and not by stock-specific quotas. Although quotas have been set in few communities, it is not known if they are adequate to prevent overhunting. A quota system for walrus hunting in Greenland began in 2006. The current control system is largely effective in ensuring the quotas are applied and that reporting is correct. Greenland currently sets quotas based on recommendations from scientific assessments using recent population estimates to allow population growth from a depleted population. A challenge with respect to managing walrus hunting remains the variable and sometimes high rates of lost animals. Since the 1960s changes in socio-economics in hunting areas of Arctic Canada and Greenland (and the use of snowmobiles instead of dog sleds in Canada) have led to a general decrease in interest in hunting of walruses and reduced harvest on walrus stocks in these countries. Although there is an active ongoing cooperation between Canada and Greenland scientists regarding assessments of shared populations of walruses currently there is no formal agreement between the two range states on co-management of shared stocks. Protection of walrus from other anthropogenic impacts generally focusses on large-scale industrial activity. The level of protection afforded to walrus habitat in many areas depends entirely on the rigor with which the Environmental Impact Assessments are conducted. Basic information on walrus such as numbers and stock discreteness is often lacking and sufficient lead-time is required to collect baseline data. Moreover, although most environmental protection legislation considers ‘cumulative impacts’, practical application remains problematic. The effectiveness of environmental protection regulations depends on industry compliance and the management authorities’ ability to enforce compliance. Because walrus are found in remote locations, enforcement remains a challenge. Increased human activity allowed by the current change in distribution and quality of arctic sea ice poses new threats to walrus if not well regulated. International agreements have varying importance for management within and among member states. Regulations governing international trade serve to identify illegally obtained products and to encourage range states to have a sustainable quota system. International cooperation in information sharing has had clear benefits for management of walruses in the past. The maintenance and expansion of these international efforts will improve the management of Atlantic walruses in the future.

Population dynamics of walruses in Greenland

The historical and current dynamics of the three Atlantic walrus (Odobenus rosmarus rosmarus) populations that occur in Greenland are estimated using age- and sex-structured population models with exponential growth, density-regulated growth and selection-delayed dynamics. These models are integrated with data in a Bayesian framework, where the likelihood of the simulated population trajectories are evaluated from recent abundance estimates and age-structure information from a selective hunt. The overall decline in the Baffin Bay population caused by historical catches is unclear due to incomplete catch reporting prior to 1950s. However, it is estimated that the population declined by 40% from the 1960s to 2005; decreased catches (≈ 140 to ≈ 70) have subsequently allowed this population to increase. The 2012 abundance estimate is 1,400 (95% CI: 1,000-2,000) individuals, and the annual natural growth rate in this population is now 7.7% (95% CI: 6.7-8.9%). Averaging across models, it is estimated that West Greenland/Baffin Island walruses declined by 80% from 7,000 (95% CI: 5,400-10,000) in 1900 to 1,350 (CI: 950-1,950) in 1960. Hereafter they increased to 3,100 (95% CI: 2,500-4,400) in 1993, and owing to increased catches they have experienced a minor decline between 1994 and the early 2000s. Annual catches where then cut from 190 to the current quota of 61, and the population is again increasing with a 2012 estimate of 3,900 (95% CI: 2,500-5,300) individuals. A 2012 estimate of 1,400 (95% CI: 700-3,100) walruses in East Greenland is recovered relative to 1888; the year prior to our first historical catches by European sealers. The historical trajectory, however, is uncertain: Density regulation estimates a relatively flat trajectory, with a maximal depletion in 1890 to 80% of the initial abundance, and a slow continuous increase to almost no current growth. A recovered population is also estimated by selection-delayed dynamics. However, this model estimates a continued increase, and a historical depletion to 2% in 1957. These results are only partially comparable with an earlier assessment for East Greenland. Updated abundance estimates for West Greenland, and modelling with age-structured data from Baffin Bay, have greatly improved our status estimates for Baffin Bay and West Greenland/Baffin Island.

Abundance of Atlantic walrus in Western Nares Strait, Baffin Bay Stock, during summer

Atlantic walruses (Odobenus rosmarus rosmarus) belonging to the Baffin Bay subpopulation occur year round in the North Water polynya (NOW) between NW Greenland and eastern Ellesmere Island (Canada). They are hunted for subsistence purposes by residents of the Qaanaaq area (NW Greenland) bordering the NOW to the east and by Canadian Inuit at the entrance to Jones Sound in Nunavut. During the open-water period NW Greenland is virtually devoid of walruses which concentrate along eastern and southern Ellesmere Island at this time of the year. To determine the abundance of walruses in the NOW area, aerial surveys were conducted in August of 1999, 2008, and 2009. In July 2009, nine satellite-linked transmitters were deployed in nearby Kane Basin. Surveys on 9 and 20 August 2009 along eastern Ellesmere Island were the most extensive and were augmented with concomitant data on haul-out and at water surface activity from three (1 F, 2 M) of the nine tags that were still functioning. We therefore focus on the 2009 surveys. Walruses were observed on the ice and in water primarily in Buchanan Bay and Princess Marie Bay where the remaining functional tags were located. The Minimum Counted population (MCP) was 571 on 20 August. Adjusting the MCP of walruses on ice for those not hauled out, the estimate of abundance of walruses in the Baffin Bay stock was 1,251(CV=1.00, 95% CI = 1,226) when adjusted by the proportion of tags ‘dry’ at the time of the survey and 1,249 (CV=1.12, 95% CI = 1,370) when adjusted by the average time tags were dry. The surveys did not cover all potential walrus summering habitat along eastern Ellesmere Island and are negatively biased to an unknown degree.