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.     

Antibodies to marine caliciviruses in the Pacific walrus (Odobenus rosmarus divergens Illiger)

Sera from 155 Pacific walruses (Odobenus rosmarus divergens Illiger), sampled in the Chukchi Sea during the summer of 1983, were tested for serum neutralizing (SN) antibodies to six marine calicivirus serotypes. Serotypes tested included San Miguel sea lion virus (SMSV) types 1, 5, 8, and 10, previously isolated from northern fur seals (Callorhinus ursinus Linné) in the Bering Sea; walrus calicivirus (WCV), previously isolated from walrus feces collected off sea ice in the Chukchi Sea; and Tillamook calicivirus (TCV), a bovine isolate from Oregon of suspected marine origin. No antibodies were found to SMSV-1, SMSV-10, or TCV. Antibodies to SMSV-5 were found in two animals (titers 1:20 and 1:160); antibodies to SMSV-8 were found in four animals (all 1:20); and antibodies to WCV were found in one animal (titer 1:40). Antibodies to WCV have been found in the Pacific walrus previously; however, this represents the first report of antibodies to any of the SMSV serotypes in this marine mammal.     

Habitat selection by ice-associated pinnipeds near St. Lawrence Island,Alaska in March 2001

Aerial surveys of ice-associated pinnipeds were conducted south of St. Lawrence Island in March 2001. The observed distributions of bearded seals (Erignathus barbatus), ribbon seals (Phoca fasciata), ringed seals (P. hispida), spotted seals (P. largha), and walruses (Odobenus rosmarus) were compared to the distributions of ice habitat types and benthic communities. Randomization tests were used to investigate habitat selection for each species. Both ringed seals and walruses preferred large ice floes (>48 m in diameter) that were common in the interior ice pack. Spotted seals favored smaller ice floes (<20 m in diameter) common near the ice edge, and bearded seals avoided large floes and preferred transitional habitat between small and large floes. Ringed seals also seemed to prefer areas with greater than 90% sea ice coverage, and bearded seals preferred 70–90% sea ice coverage while avoiding areas with greater than 90% coverage. All species, except spotted seals, were seen most frequently in a region of high benthic biomass, and randomization tests suggested that bearded seals actively selected that region.     

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.     

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.     

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

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

Underwater observations of foraging free-living Atlantic walruses (Odobenus rosmarus rosmarus) and estimates of their food consumption

Food consumption of Atlantic walruses (Odobenus rosmarus rosmarus L.) was quantified by combining underwater observations of feeding with satellite-telemetry data on movement and diving activity. The study was conducted between 31 July and 7 August 2001 in Young Sound (74°N-20°W) in Northeast Greenland. On ten occasions, divers were able to accompany foraging walruses to the sea floor and collect the shells of newly predated bivalves (Mya truncata, Hiatella arctica, Serripes groenlandicus) for determination of number of prey and biomass ingested per dive. Simultaneously, the activity of a 1,200-kg adult male walrus was studied by use of satellite-telemetry during an entire foraging cycle that included 74 h at sea followed by a 23-h rest on land. An average of 53.2 bivalves (SE=5.2, range: 34-89, n=10) were consumed per dive, corresponding to 149.0 g shell-free dry matter (SE=18.9, range: 62.4-253.1 g), or 2,576 kJ per dive (SE=325.2, range: 1,072-4,377 kJ). During the foraging trip, the walrus spent 57% of the time diving to depths of between 6 and 32 m, and it made a total of 412 dives that lasted between 5 and 7 min (i.e. typical foraging dives). If the entire feeding cycle is considered (97 h), the estimated daily gross energy intake was 214 kJ per kg body mass (95% CI: 153-275 kJ), corresponding to the ingestion of 57 kg (95% CI: 41-72 kg) wet weight bivalve biomass per day, or 4.7 (95% CI: 3.3-5.9%) of total walrus body mass. Due to ice cover, walrus access to the plentiful inshore bivalve banks in the area is restricted to the short summer period, where walruses rely on them for replenishing energy stores. It is hypothesised that the documented decrease in the extent and duration of Arctic sea ice may increase food availability for walruses in eastern Greenland in the future.     

A temporal comparison of a benthic infaunal community southwest of St. Lawrence Island, Bering Sea between 2006 and 1970–1974

Benthic infaunal biomass and abundance may be changing in Bering Sea communities. This study compared benthic infaunal biomass, abundance, and community composition southwest of St. Lawrence Island in an important forage area for benthic-feeding birds and mammals between 1970–1974 and 2006. Invertebrate biomass and abundance were significantly greater in 2006 than in 1970–1974 primarily due to high nuculid (Bivalvia) biomass and abundance that contributed 13.2% (biomass) and 8.5% (abundance) to differences in community structure between the sampling periods. This is in contrast to St. Lawrence Island Polynya studies conducted in the 1980s and 1990s that documented decreases in benthic biomass and abundance. Spatial scale of sampling, selective predation, a strong recruitment event, and/or sampling design may account for the difference in trends among the studies.     

Large numbers of marine mammals winter in the North Water polynya

The importance of the North Water polynya in Smith Sound as an overwintering area for marine mammals has been questioned. One way to address the issue is to assess the abundance of selected marine mammals that are present during winter in the North Water. Visual aerial surveys involving double observer platforms were conducted over the eastern part of the North Water polynya in April 2014. Four species of marine mammals were included in strip-census estimation of abundance. Perception bias was addressed using a double-platform survey protocol, a Chapman mark–recapture estimator for whales, seals and walruses (Odobenus rosmarus) on ice and a mark–recapture distance sampling estimation technique for walruses in water. Availability bias was addressed by correcting abundance estimates by the percentage of time animals detected in water that were available for detection at the surface. The resulting estimates suggested that 2544 walruses (95 % CI 1513–4279), 6005 bearded seals (Erignathus barbatus, 95 % CI 4070–8858), 2324 belugas (Delphinapterus leucas, 95 % CI 968–5575) and 3059 narwhals (Monodon monoceros, 95 % CI 1760–5316) wintered in the eastern part of the North Water polynya in April 2014. The walrus estimate is larger than previous summer estimates, and it emphasizes the importance of the habitat along the Greenland coast as a walrus wintering ground. The estimate of belugas is likely negatively biased due to the partial coverage of the potential habitat. The estimate of narwhals is large compared to the few previous observations of narwhals in winter in the North Water, and it demonstrates that large numbers of narwhals winter there. The overall conclusion is that the North Water is indeed an important wintering area for at least walruses, belugas, narwhals and bearded seals.     

A morbillivirus antibody survey of Atlantic walrus, narwhal and beluga in Canada

A longitudinal serologic survey was conducted for morbillivirus antibodies in Atlantic walruses (Odobenus rosmarus rosmarus), narwhal (Monodon monoceros), and beluga (Delphinapterus leucas) from the Northwest Territories, Nunavut and the St. Lawrence estuary (Canada). Sixty-five of 131 (50%) walruses sampled between 1984 and 1993 had detectable morbillivirus neutralizing antibodies. Positive walrus were identified from four of five Arctic sampling sites, to as far back as 1984. Prevalence of morbillivirus neutralizing antibodies in walruses from Foxe Basin ranged from a high of 76% (n = 21) in 1993 to a low of 22% (n = 28) in 1984. Limitations in sample acquisition may have produced underestimates for the 1984 data. There are no reports of clinical morbillivirus infection in walruses. Our results are consistent with the hypothesis that a morbillivirus similar or identical to phocine distemper virus (PDV) has circulated among walrus populations of the eastern Canadian Arctic, at least since the early 1980s. No narwhal (n = 79) or beluga (n = 445) from Arctic waters were identified as having antibodies to dolphin morbilivirus (DMV) above the threshold serum dilution of log2 4. Also, none of the beach-cast cetacean carcasses (n = 28) from the Gulf of St. Lawrence and the St. Lawrence estuary were positive for antibodies to DMV. This indicates that Gulf of St. Lawrence, St. Lawrence estuary, and Arctic cetaceans either have not been exposed to DMV or an antigenically related morbillivirus, or are not susceptible to infection.     

Inter-annual variability in the proportional contribution of higher trophic levels to the diet of Pacific walruses

Pacific walruses (Odobenus rosmarus divergens) depend on Arctic sea ice as a resting and foraging platform; however, recent years have seen unprecedented seasonal reductions in ice extent. Previous researchers proposed that during unfavorable ice conditions, walruses might prey on other pinnipeds. To examine this hypothesis, we analyzed carbon and nitrogen stable isotope ratios of muscle from walruses (n = 155) sampled from the Bering and Chukchi seas during 2001–2010. We used a Bayesian stable isotope mixing model to examine the proportional contribution of higher trophic level prey (HTLP) (e.g., seals, seabirds) to walrus diets and extrapolated a tissue-specific turnover rate to compare diet of individuals over time. Mode HTLP across years was 19 % ± 8. Results indicate a significant decrease (P < 0.05) in the reliance on HTLP during 2008–2009 (mode HTLP 13 %), one of two sampling periods that experienced great seasonal loss of pan-Arctic sea ice (the other being 2007–2008 with mode HTLP of 23 %). We also reveal intra-annual fluctuations in the contribution of HTLP to the diet of a walrus sampled in 2011 with seal remains in its stomach through high-resolution sectioning along a whisker length. Our findings suggest that walruses forage opportunistically as a result of multiple environmental factors and that sea ice extent alone does not drive consumption of HTLP.     

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.     

Population structure of North American beluga whales (Delphinapterus leucas) based on nuclear DNA microsatellite variation and contrasted with the population structure revealed by mitochondrial DNA variation

Beluga whales ( Delphinapterus leucas ) in North American waters migrate seasonally between wintering areas in broken pack ice and summering locations in estuaries and other open water areas in the Arctic and sub-Arctic. Results from our previous investigation of beluga whale mitochondrial DNA (mtDNA) revealed genetic heterogeneity among beluga from different summering locations that was interpreted as representing a high degree of summering site philopatry. However, mtDNA is maternally inherited and does not reflect mating that may occur among beluga from different summering locations in wintering areas or during annual migrations. To test the possibility that breeding occurs among beluga from different summering locations, genetic variability at five nuclear DNA (nDNA) microsatellite loci was examined in the same animals tested in the mtDNA study. Beluga samples ( n = 640) were collected between 1984 and 1994 from 24 sites across North America, mostly during the summer. Whales from the various sites were categorized into eight summering locations as identified by mtDNA analysis, as well as four hypothesized wintering areas: Bering Sea, Hudson Strait (Hudson Strait, Labrador Sea, southwest Davis Strait), Baffin Bay (North Water, east Davis Strait), and St Lawrence River. Microsatellite allele frequencies indicated genetic homogeneity among animals from summering sites believed to winter together but differentiation among whales from some of the wintering areas. In particular, beluga from western North America (Bering Sea) were clearly distinguished from beluga from eastern North America (Hudson Strait, Baffin Bay, and St Lawrence River). Based upon the combined data set, the population of North American beluga whales was divided into two evolutionarily significant units. However, the population may be further subdivided into management units to reflect distinct groups of beluga at summering locations.     

Population structure of North American beluga whales (Delphinapterus leucas) based on nuclear DNA microsatellite variation and contrasted with the population structure revealed by mitochondrial DNA variation

Beluga whales ( Delphinapterus leucas ) in North American waters migrate seasonally between wintering areas in broken pack ice and summering locations in estuaries and other open water areas in the Arctic and sub-Arctic. Results from our previous investigation of beluga whale mitochondrial DNA (mtDNA) revealed genetic heterogeneity among beluga from different summering locations that was interpreted as representing a high degree of summering site philopatry. However, mtDNA is maternally inherited and does not reflect mating that may occur among beluga from different summering locations in wintering areas or during annual migrations. To test the possibility that breeding occurs among beluga from different summering locations, genetic variability at five nuclear DNA (nDNA) microsatellite loci was examined in the same animals tested in the mtDNA study. Beluga samples ( n = 640) were collected between 1984 and 1994 from 24 sites across North America, mostly during the summer. Whales from the various sites were categorized into eight summering locations as identified by mtDNA analysis, as well as four hypothesized wintering areas: Bering Sea, Hudson Strait (Hudson Strait, Labrador Sea, southwest Davis Strait), Baffin Bay (North Water, east Davis Strait), and St Lawrence River. Microsatellite allele frequencies indicated genetic homogeneity among animals from summering sites believed to winter together but differentiation among whales from some of the wintering areas. In particular, beluga from western North America (Bering Sea) were clearly distinguished from beluga from eastern North America (Hudson Strait, Baffin Bay, and St Lawrence River). Based upon the combined data set, the population of North American beluga whales was divided into two evolutionarily significant units. However, the population may be further subdivided into management units to reflect distinct groups of beluga at summering locations.     

Seasonal variation in Arctic marine mammal acoustic detection in the northern Bering Sea

Declines in Arctic sea ice cover are influencing the distribution of protected endemic marine mammals, many of which are important for local Indigenous Peoples, and increasing the presence of potentially disruptive industrial activities. Due to increasing conservation concerns, we conducted the first year-round acoustic monitoring of waters off Gambell and Savoonga (St. Lawrence Island, Alaska), and in the Bering Strait to quantify vocalizing presence of bowhead whales, belugas, walruses, bearded seals, and ribbon seals. Bottom-mounted archival acoustic recorders collected data for up to 10 months per deployment between 2012 and 2016. Spectrograms were analyzed for species-typical vocalizations, and daily detection rates and presence/absence were calculated. Generalized additive models were used to model call presence as a function of time-of-year, sea surface temperature, and sea ice concentration. We identified seasonality in call presence for all species, corroborating previous acoustic and distribution studies, and identified finer-scale spatiotemporal distribution via occurrence of call presence between different monitoring sites. Time-of-year was the strongest significant effect on call presence for all species. These data provide important information on Arctic endemic species' spatiotemporal distributions in biologically and culturally important areas within a rapidly changing Arctic region.     

Pacific walrus: Benthic bioturbator of Beringia

The dependency of walruses on sea ice as habitat, the extent of their feeding, their benthic bioturbation and consequent nutrient flux suggest that walruses play a major ecological role in Beringia. This suggestion is supported by several lines of evidence, accumulated during more than three decades of enquiry and leading to the hypothesis that positive feedbacks of walrus feeding strongly influence productivity and ecological function via benthic bioturbation and nutrient flux. Walruses annually consume an estimated 3 million metric tons of benthic biomass. Walrus prey species inhabit patches across the shelf according to sediment type and structure. Side-scan sonar and our calculations indicate that the area affected by walrus feeding is in the order of thousands of square kilometers per year. Annual to long-term walrus bioturbation results in significant, large-scale changes in sediment and biological-community structure, and magnifies nutrient flux from sediment pore water to the water column by about two orders of magnitude over wide areas. The combined effects of walrus feeding must be placed in the context of long-term, regional climate changes and responses. Should sea ice continue to move northward as a result of climate change, the walrus' ecological role could be diminished or lost, the benthic ecosystem could be fundamentally altered and native subsistence hunters would be deprived of important resources.     

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.     

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.     

Faecal DNA amplification in Pacific walruses (Odobenus rosmarus divergens)

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

Pacific Walrus and climate change: observations and predictions

The extent and duration of sea-ice habitats used by Pacific walrus (Odobenus rosmarus divergens) are diminishing resulting in altered walrus behavior, mortality, and distribution. I document changes that have occurred over the past several decades and make predictions to the end of the 21st century. Climate models project that sea ice will monotonically decline resulting in more ice-free summers of longer duration. Several stressors that may impact walruses are directly influenced by sea ice. How these stressors materialize were modeled as most likely-case, worst-case, and best-case scenarios for the mid- and late-21st century, resulting in four comprehensive working hypotheses that can help identify and prioritize management and research projects, identify comprehensive mitigation actions, and guide monitoring programs to track future developments and adjust programs as needed. In the short term, the most plausible hypotheses predict a continuing northward shift in walrus distribution, increasing use of coastal haulouts in summer and fall, and a population reduction set by the carrying capacity of the near shore environment and subsistence hunting. Alternatively, under worst-case conditions, the population will decline to a level where the probability of extinction is high. In the long term, walrus may seasonally abandon the Bering and Chukchi Seas for sea-ice refugia to the northwest and northeast, ocean warming and pH decline alter walrus food resources, and subsistence hunting exacerbates a large population decline. However, conditions that reverse current trends in sea ice loss cannot be ruled out. Which hypothesis comes to fruition depends on how the stressors develop and the success of mitigation measures. Best-case scenarios indicate that successful mitigation of unsustainable harvests and terrestrial haulout-related mortalities can be effective. Management and research should focus on monitoring, elucidating effects, and mitigation, while ultimately, reductions in greenhouse gas emissions are needed to reduce sea-ice habitat losses.