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 0-2-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(-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(-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's body mass over the first month after parturition if none of the lactation costs is met through ingested prey.

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 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: 0.92-1.68 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 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.

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(-1) for 2-yr-olds to 93,370 kcal d(-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.

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.

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.