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Chadwick V. Jay

Publications and source records attributed to Chadwick V. Jay.

At least 19 recordsLinked to original sources

Exploring effects of vessels on walrus behaviors using telemetry, automatic identification system data and matching

Arctic marine mammals have had little exposure to vessel traffic and potential associated disturbance, but sea ice loss has increased accessibility of Arctic waters to vessels. Vessel disturbance could influence marine mammal population dynamics by altering behavioral activity budgets that affect energy balance, which in turn can affect birth and death rates. As an initial step in studying these linkages, we conducted the first comprehensive analysis to evaluate the effects of vessel exposure on Pacific walrus ( Odobenus rosmarus divergens ) behaviors. We obtained >120,000 h of location and behavior (foraging, in-water not foraging, and hauled out) data from 218 satellite-tagged walruses and linked them to vessel locations from the marine automatic identification system (AIS). This yielded 206 vessel-exposed walrus telemetry hours for comparison to unexposed hours, which we used to assess if vessel exposure altered walrus behavior. We developed a filter to account for misclassification of vessel exposure of telemetered walruses. Then we tested for an effect of vessel exposure on walrus behaviors using a combination of exact and propensity score-based matching to account for confounding covariates, and we conducted statistical power analyses. We did not detect an effect of vessel exposure on walrus behaviors even when statistical power was high (i.e., for foraging walruses), which may have been due to the sample size-driven need to define vessel presence within a larger than desired distance (15-km measured radius) around a walrus. Although this study did not determine at what distance vessel exposure affects walrus behaviors, it provided an upper bound on the distance at which the vessels encountered may disturb foraging walruses. When more situation-specific information is lacking, this distance could be used as a conservative buffer to maintain between vessels and areas of high use by foraging walruses. Studies on behavioral consequences of closer proximities between walruses and vessels are needed, and our assessments of misclassification rates and statistical power can be used for future studies. We demonstrated that analytical approaches such as matching, which are rarely used in wildlife studies, are particularly useful for testing hypotheses with observational data.

Alaska

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.

Alaska

Variability of lipids and fatty acids in Pacific walrus blubber

The variability of lipid content and fatty acid (FA) composition across blubber depth and body sites are important considerations for condition and diet studies of marine mammals. We investigated lipid and FA variability among inner and outer blubber layers, three body sites, four study years, and lactation status of adult female Pacific walruses ( Odobenus rosmarus divergens ) using blubber samples collected from subsistence-harvested walruses in spring 2007–2010. Percent lipid content did not differ between the inner and outer blubber layers at the rump, flank, or sternum of walruses. Although FA composition differed between the inner and outer blubber layers, the difference was consistent across body sites, and differences between layers within individual FAs were small (<2%). Lipid content at the sternum of lactating females was 6% higher than non-lactating females, consistent with known variation in body condition among these reproductive classes. Across study years, lipid content varied 18% and individual FAs varied 6%, likely reflecting population-level interannual variability in energy budgets and small differences in diet among years. Consistency in blubber lipid content across blubber depth and body sites and detectable variation in blubber lipid content among reproductive classes and years suggests the potential for lipid content to be a useful indicator of walrus body condition. In addition to information on condition, FA composition of blubber samples could potentially provide insights into changes in walrus diet that may be expected to occur from changes in their access to prey resources resulting from continued sea ice loss.

Frontiers in Marine Science

DNA metabarcoding of feces to infer summer diet of Pacific walruses

Environmental conditions in the Chukchi Sea are changing rapidly and may alter the abundance and distribution of marine species and their benthic prey. We used a metabarcoding approach to identify potentially important prey taxa from Pacific walrus ( Odobenus rosmarus divergens ) fecal samples ( n = 87). Bivalvia was the most dominant class of prey (66% of all normalized counts) and occurred in 98% of the samples. Polychaeta and Gastropoda occurred in 70% and 62% of the samples, respectively. The remaining nine invertebrate classes comprised <21% of all normalized counts. The common occurrence of these three prey classes is consistent with examinations of walrus stomach contents. Despite these consistencies, biases in the metabarcoding approach to determine diet from feces have been highlighted in other studies and require further study, in addition to biases that may have arisen from our opportunistic sampling. However, this noninvasive approach provides accurate identification of prey taxa from degraded samples and could yield much‐needed information on shifts in walrus diet in a rapidly changing Arctic.

Alaska

A multi-species synthesis of satellite telemetry data in the Pacific Arctic (1987–2015): Overlap of marine mammal distributions and core use areas

We collated available satellite telemetry data for six species of ice-associated marine mammals in the Pacific Arctic: ringed seals ( Pusa hispida ; n = 118), bearded seals( Erignathus barbatus, n = 51), spotted seals ( Phoca largha , n = 72), Pacific walruses ( Odobenus rosmarus divergens, n = 389); bowhead whales ( Balaena mysticetus , n = 46), and five Arctic and sub-arctic stocks of beluga whales ( Delphinapterus leucas , n = 103). We also included one seasonal resident, eastern North Pacific gray whales ( Eschrichtius robustus , n = 12). This review summarized the distribution of daily locations from satellite-linked transmitters during two analysis periods, summer (May–November) and winter (December–April), and then examined the overlap among species. Six multi-species core use areas were identified during the summer period: 1) Chukotka/Bering Strait; 2) Norton Sound; 3) Kotzebue Sound; 4) the northeastern Chukchi Sea; 5) Mackenzie River Delta/Amundsen Gulf; and 6) Viscount Melville Sound. During the winter period, we identified four multi-species core use areas: 1) Anadyr Gulf/Strait; 2) central Bering Sea; 3) Nunivak Island; and 4) Bristol Bay. During the summer period, four of the six areas were centered on the greater Bering Strait region and the northwestern coast of Alaska and included most of the species we examined. The two remaining summer areas were in the western Canadian Arctic and were largely defined by the seasonal presence of Bering-Chukchi-Beaufort stock bowhead whales and Eastern Beaufort Sea stock beluga whales, whose distribution overlapped during both summer and winter periods. During the winter period, the main multi-species core use area was located near the Gulf of Anadyr and extended northwards through Anadyr and Bering Straits. This area is contained within the Bering Sea “green belt”, an area of enhanced primary and secondary productivity in the Bering Sea. We also described available telemetry data and where they can be found as of 2017. These data are important for understanding ice-associated marine mammal movements and habitat use in the Pacific Arctic and should be archived, with appropriate metadata, to ensure they are available for future retrospective analyses.

Deep Sea Research Part II: Topical Studies in Ocea

Demography of the Pacific walrus (Odobenus rosmarus divergens) in a changing Arctic

The Pacific walrus ( Odobenus rosmarus divergens ) is a candidate to be listed as an endangered species under United States law, in part, because of climate change‐related concerns. While the population was known to be declining in the 1980s and 1990s, its recent status has not been determined. We developed Bayesian models of walrus population dynamics to assess the population by synthesizing information on population sizes, age structures, reproductive rates, and harvests for 1974–2015. Candidate models allowed for temporal variation in some or all vital rates, as well as density dependence or density independence in reproduction and calf survival. All selected models indicated that the population underwent a multidecade decline, which began moderating in the 1990s, and that annual reproductive rate and natural calf survival rates rose over time in a density‐dependent manner. However, selected models were equivocal regarding whether the natural juvenile survival rate was constant or decreasing over time. Depending on whether juvenile survival decreased after 1998, the population growth rate either increased during 1999–2015 or stabilized at a lesser level of decline than seen in the 1980s. The probability that the population was still declining in 2015 ranged from 45% to 87%.

Marine Mammal Science

Walrus haul-out and in water activity levels relative to sea ice availability in the Chukchi Sea

An animal’s energetic costs are dependent on the amount of time it allocates to various behavioral activities. For Arctic pinnipeds, the time allocated to active and resting behaviors could change with future reductions in sea ice cover and longer periods of open water. The Pacific walrus ( Odobenus rosmarus divergens ) is a large Arctic pinniped that rests on sea ice or land between foraging trips to feed on the seafloor. We used behavioral data collected from radiotagged walruses in the Chukchi Sea (2008–2014) in a Bayesian generalized linear mixed effects model to estimate the probability a walrus was in water foraging, in water not foraging, or hauled out, as a function of environmental covariates. The probability of a walrus being in water increased with wind speed and decreased with air temperature, and the probability a walrus was foraging, given it was in water, increased with available benthic macrofaunal biomass. The probability of each behavior was also related to the nature and availability of haul-out substrates. The amount of time walruses spent in water foraging and hauled out was greatest when only sea ice was available, which typically occurs when walruses occupy feeding areas during summer and early autumn. This situation may be most energy efficient for walruses because it allows the highest proportion of in water energy expenditure to be allocated to foraging. Conversely, the amount of time walruses spent in water foraging and hauled out was lowest when only land was available, which typically occurs in late autumn, in years when walruses were constrained to land haul-outs because sea ice was absent over the continental shelf.

Chukchi Sea

Evaluation of a method using survey counts and tag data to estimate the number of Pacific walruses ( Odobenus rosmarus divergens ) using a coastal haulout in northwestern Alaska

Increased periods of sparse sea ice over the continental shelf of the Chukchi Sea in late summer have reduced offshore haulout habitat for Pacific walruses ( Odobenus rosmarus divergens ) and increased opportunities for human activities in the region. Knowing how many walruses could be affected by human activities would be useful to conservation decisions. Currently, there are no adequate estimates of walrus abundance in the northeastern Chukchi Sea during summer–early autumn. Estimating abundance in autumn might be possible from coastal surveys of hauled out walruses during periods when offshore sea ice is unavailable to walruses. We evaluated methods to estimate the size of the walrus population that was using a haulout on the coast of northwestern Alaska in autumn by using aerial photography to count the number of hauled out walruses (herd size) and data from 37 tagged walruses to estimate availability (proportion of population hauled out). We used two methods to estimate availability, direct proportions of hauled out tagged walruses and smoothed proportions using local polynomial regression. Point estimates of herd size (4200–38,000 walruses) and total population size (76,000–287,000 walruses) ranged widely among days and between the two methods of estimating availability. Estimates of population size were influenced most by variation in estimates of availability. Coastal surveys might be improved most by counting walruses when the greatest numbers are hauled out, thereby reducing the influence of availability on population size estimates. The chance of collecting data during peak haulout periods would be improved by conducting multiple surveys.

Polar Biology

A strategy for recovering continuous behavioral telemetry data from Pacific walruses

Tracking animal behavior and movement with telemetry sensors can offer substantial insights required for conservation. Yet, the value of data collected by animal-borne telemetry systems is limited by bandwidth constraints. To understand the response of Pacific walruses ( Odobenus rosmarus divergens ) to rapid changes in sea ice availability, we required continuous geospatial chronologies of foraging behavior. Satellite telemetry offered the only practical means to systematically collect such data; however, data transmission constraints of satellite data-collection systems limited the data volume that could be acquired. Although algorithms exist for reducing sensor data volumes for efficient transmission, none could meet our requirements. Consequently, we developed an algorithm for classifying hourly foraging behavior status aboard a tag with limited processing power. We found a 98% correspondence of our algorithm's classification with a test classification based on time–depth data recovered and characterized through multivariate analysis in a separate study. We then applied our algorithm within a telemetry system that relied on remotely deployed satellite tags. Data collected by these tags from Pacific walruses across their range during 2007–2015 demonstrated the consistency of foraging behavior collected by this strategy with data collected by data logging tags; and demonstrated the ability to collect geospatial behavioral chronologies with minimal missing data where recovery of data logging tags is precluded. Our strategy for developing a telemetry system may be applicable to any study requiring intelligent algorithms to continuously monitor behavior, and then compress those data into meaningful information that can be efficiently transmitted.

Wildlife Society Bulletin

An evaluation of behavior inferences from Bayesian state-space models: A case study with the Pacific walrus

State-space models offer researchers an objective approach to modeling complex animal location data sets, and state-space model behavior classifications are often assumed to have a link to animal behavior. In this study, we evaluated the behavioral classification accuracy of a Bayesian state-space model in Pacific walruses using Argos satellite tags with sensors to detect animal behavior in real time. We fit a two-state discrete-time continuous-space Bayesian state-space model to data from 306 Pacific walruses tagged in the Chukchi Sea. We matched predicted locations and behaviors from the state-space model (resident, transient behavior) to true animal behavior (foraging, swimming, hauled out) and evaluated classification accuracy with kappa statistics ( κ ) and root mean square error (RMSE). In addition, we compared biased random bridge utilization distributions generated with resident behavior locations to true foraging behavior locations to evaluate differences in space use patterns. Results indicated that the two-state model fairly classified true animal behavior (0.06 ≤ κ ≤ 0.26, 0.49 ≤ RMSE ≤ 0.59). Kernel overlap metrics indicated utilization distributions generated with resident behavior locations were generally smaller than utilization distributions generated with true foraging behavior locations. Consequently, we encourage researchers to carefully examine parameters and priors associated with behaviors in state-space models, and reconcile these parameters with the study species and its expected behaviors.

Marine Mammal Science

Sex-specific energetics of Pacific walruses ( Odobenus rosmarus divergens ) during the nursing interval

Habitat use and activity patterns of Pacific walruses ( Odobenus rosmarus divergens ) have changed with climate-induced reductions in sea ice. Increases in the time active in water could result in negative energy balance, precluding females from sustaining lactation, which could impact population demographics. Little is known about lactation costs in walruses. We examined the energetics of 0&ndash;2-yr-old walrus calves by using Bayesian hierarchical models based on longitudinal husbandry records of growth ( n = 6 females and 7 males) and caloric intake ( n = 5 females and 6 males) as a proxy for maternal lactation costs. Males and females had similar growth patterns; mean mass increased from 68 kg at birth to 301 kg by 2 yr. Females had a 2,000 kcal kg &minus;1 higher mass storage (growth) cost than males; females typically synthesize and deposit greater amounts of adipose, which is more energy dense than lean tissue. In contrast, males had higher metabolic (basal and activity) costs, ranging from 600 to 1,800 kcal d &minus;1 greater than similarly sized females; males are typically leaner, and muscle is more metabolically active than adipose. Yet total daily energy requirements (storage plus metabolic components) were similar across sexes, summing to approximately 190,000 kcal over the first month postpartum. Based on these estimates and assuming that 8,103 kcal is recovered from 1 kg of mass loss in adult female walruses, suckling calves could deplete 23 kg of their mother&rsquo;s body mass over the first month after parturition if none of the lactation costs is met through ingested prey.

Physiological and Biochemical Zoology

Pacific walrus coastal haulout database, 1852-2016— Background report

Walruses are large benthic predators that rest out of water between foraging bouts. Coastal &ldquo;haulouts&rdquo; (places where walruses rest) are formed by adult males in summer and sometimes by females and young when sea ice is absent, and are often used repeatedly across seasons and years. Understanding the geography and historical use of haulouts provides a context for conservation efforts. We summarize information on Pacific walrus haulouts from available reports (n =151), interviews with coastal residents and aviators, and personal observations of the authors. We provide this in the form of a georeferenced database that can be queried and displayed with standard geographic information system and database management software. The database contains 150 records of Pacific walrus haulouts, with a summary of basic characteristics on maximum haulout aggregation size, age-sex composition, season of use, and decade of most recent use. Citations to reports are provided in the appendix and as a bibliographic database. Haulouts were distributed across the coasts of the Pacific walrus range; however, the largest (maximum >10,000 walruses) of the haulouts reported in the recent 4 decades (n=19) were concentrated on the Russian shores in regions near the Bering Strait and northward into the western Chukchi Sea (n=17). Haulouts of adult female and young walruses primarily occurred in the Bering Strait region and areas northward, with others occurring in the central Bering Sea, Gulf of Anadyr, and Saint Lawrence Island regions. The Gulf of Anadyr was the only region to contain female and young walrus haulouts, which formed after the northward spring migration and prior to autumn ice formation.

Alaska

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

Physiological constraints dictate animals&rsquo; 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 3 month old and 20 adults), ranging from 41.31 to 54.14 slykes and 42.00 to 46.93 slykes in the longissimus and supraspinatus, respectively. Mean myoglobin in the fetus and neonatal calves fell within a narrow range (longissimus: 0.92&ndash;1.68 g 100 g &minus;1 wet muscle mass; supraspinatus: 0.88&ndash;1.64 g 100 g &minus;1 wet muscle mass). By 3 months post-partum, myoglobin in the longissimus increased by 79%, but levels in the supraspinatus remained unaltered. From 3 months 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.

Journal of Experimental Biology

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 &times; standard length &minus;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&ndash;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.

Marine Mammal Science

Pacific walrus ( Odobenus rosmarus divergens ) resource selection in the northern Bering Sea

The Pacific walrus is a large benthivore with an annual range extending across the continental shelves of the Bering and Chukchi Seas. We used a discrete choice model to estimate site selection by adult radio-tagged walruses relative to the availability of the caloric biomass of benthic infauna and sea ice concentration in a prominent walrus wintering area in the northern Bering Sea (St. Lawrence Island polynya) in 2006, 2008, and 2009. At least 60% of the total caloric biomass of dominant macroinfauna in the study area was composed of members of the bivalve families Nuculidae, Tellinidae, and Nuculanidae. Model estimates indicated walrus site selection was related most strongly to tellinid bivalve caloric biomass distribution and that walruses selected lower ice concentrations from the mostly high ice concentrations that were available to them (quartiles: 76%, 93%, and 99%). Areas with high average predicted walrus site selection generally coincided with areas of high organic carbon input identified in other studies. Projected decreases in sea ice in the St. Lawrence Island polynya and the potential for a concomitant decline of bivalves in the region could result in a northward shift in the wintering grounds of walruses in the northern Bering Sea.

PLoS ONE

Spatial variation and low diversity in the major histocompatibility complex in walrus ( Odobenus rosmarus )

Increased global temperature and associated changes to Arctic habitats will likely result in the northward advance of species, including an influx of pathogens novel to the Arctic. How species respond to these immunological challenges will depend in part on the adaptive potential of their immune response system. We compared levels of genetic diversity at a gene associated with adaptive immune response [Class II major histocompatibility complex (MHC), DQB exon 2] between populations of walrus ( Odobenus rosmarus ), a sea ice-dependent Arctic species. Walrus was represented by only five MHC DQB alleles, with frequency differences observed between Pacific and Atlantic populations. MHC DQB alleles appear to be under balancing selection, and most (80 %; n = 4/5) of the alleles were observed in walruses from both oceans, suggesting broad scale differences in the frequency of exposure and diversity of pathogens may be influencing levels of heterozygosity at DQB in walruses. Limited genetic diversity at MHC, however, suggests that walrus may have a reduced capacity to respond to novel immunological challenges associated with shifts in ecological communities and environmental stressors predicted for changing climates. This is particularly pertinent for walrus, since reductions in summer sea ice may facilitate both northward expansion of marine species and associated pathogens from more temperate regions, and exchange of marine mammals and associated pathogens through the recently opened Northwest Passage between the Atlantic and Pacific Oceans in the Canadian high Arctic.

Polar Biology

Energy demands for maintenance, growth, pregnancy, and lactation of female Pacific walruses ( Odobenus rosmarus divergens )

Decreases in sea ice have altered habitat use and activity patterns of female Pacific walruses Odobenus rosmarus divergens and could affect their energetic demands, reproductive success, and population status. However, a lack of physiological data from walruses has hampered efforts to develop the bioenergetics models required for fully understanding potential population-level impacts. We analyzed long-term longitudinal data sets of caloric consumption and body mass from nine female Pacific walruses housed at six aquaria using a hierarchical Bayesian approach to quantify relative energetic demands for maintenance, growth, pregnancy, and lactation. By examining body mass fluctuations in response to food consumption, the model explicitly uncoupled caloric demand from caloric intake. This is important for pinnipeds because they sequester and deplete large quantities of lipids throughout their lifetimes. Model outputs were scaled to account for activity levels typical of free-ranging Pacific walruses, averaging 83% of the time active in water and 17% of the time hauled-out resting. Estimated caloric requirements ranged from 26,900 kcal d&minus;1 for 2-yr-olds to 93,370 kcal d&minus;1 for simultaneously lactating and pregnant walruses. Daily consumption requirements were higher for pregnancy than lactation, reflecting energetic demands of increasing body size and lipid deposition during pregnancy. Although walruses forage during lactation, fat sequestered during pregnancy sustained 27% of caloric requirements during the first month of lactation, suggesting that walruses use a mixed strategy of capital and income breeding. Ultimately, this model will aid in our understanding of the energetic and population consequences of sea ice loss.

Physiological and Biochemical Zoology

Estimating age ratios and size of Pacific walrus herds on coastal haulouts using video imaging

During Arctic summers, sea ice provides resting habitat for Pacific walruses as it drifts over foraging areas in the eastern Chukchi Sea. Climate-driven reductions in sea ice have recently created ice-free conditions in the Chukchi Sea by late summer causing walruses to rest at coastal haulouts along the Chukotka and Alaska coasts, which provides an opportunity to study walruses at relatively accessible locations. Walrus age can be determined from the ratio of tusk length to snout dimensions. We evaluated use of images obtained from a gyro-stabilized video system mounted on a helicopter flying at high altitudes (to avoid disturbance) to classify the sex and age of walruses hauled out on Alaska beaches in 2010–2011. We were able to classify 95% of randomly selected individuals to either an 8- or 3-category age class, and we found measurement-based age classifications were more repeatable than visual classifications when using images presenting the correct head profile. Herd density at coastal haulouts averaged 0.88 walruses/m 2 (std. err. = 0.02), herd size ranged from 8,300 to 19,400 (CV 0.03–0.06) and we documented ~30,000 animals along ~1 km of beach in 2011. Within the herds, dependent walruses (0–2 yr-olds) tended to be located closer to water, and this tendency became more pronounced as the herd spent more time on the beach. Therefore, unbiased estimation of herd age-ratios will require a sampling design that allows for spatial and temporal structuring. In addition, randomly sampling walruses available at the edge of the herd for other purposes (e.g., tagging, biopsying) will not sample walruses with an age structure representative of the herd. Sea ice losses are projected to continue, and population age structure data collected with aerial videography at coastal haulouts may provide demographic information vital to ongoing efforts to understand effects of climate change on this species.

PLoS ONE