Female Pacific walruses (Odobenus rosmarus divergens) show greater partitioning of sea ice organic carbon than males: Evidence from ice algae trophic markers.

The expected reduction of ice algae with declining sea ice may prove to be detrimental to the Pacific Arctic ecosystem. Benthic organisms that rely on sea ice organic carbon (iPOC) sustain benthic predators such as the Pacific walrus (Odobenus rosmarus divergens). The ability to track the trophic transfer of iPOC is critical to understanding its value in the food web, but prior methods have lacked the required source specificity. We analyzed the H-Print index, based on biomarkers of ice algae versus phytoplankton contributions to organic carbon in marine predators, in Pacific walrus livers collected in 2012, 2014 and 2016 from the Northern Bering Sea (NBS) and Chukchi Sea. We paired these measurements with stable nitrogen isotopes (δ15N) to estimate trophic position. We observed differences in the contribution of iPOC in Pacific walrus diet between regions, sexes, and age classes. Specifically, the contribution of iPOC to the diet of Pacific walruses was higher in the Chukchi Sea (52%) compared to the NBS (30%). This regional difference is consistent with longer annual sea ice persistence in the Chukchi Sea. Within the NBS, the contribution of iPOC to walrus spring diet was higher in females (~45%) compared to males (~30%) for each year (p < 0.001), likely due to specific foraging behavior of females to support energetic demands associated with pregnancy and lactation. Within the Chukchi Sea, the iPOC contribution was similar between males and females, yet higher in juveniles than in adults. Despite differences in the origin of organic carbon fueling the system (sea ice versus pelagic derived carbon), the trophic position of adult female Pacific walruses was similar between the NBS and Chukchi Sea (3.2 and 3.5, respectively), supporting similar diets (i.e. clams). Given the higher quality of organic carbon from ice algae, the retreat of seasonal sea ice in recent decades may create an additional vulnerability for female and juvenile Pacific walruses and should be considered in management of the species.

Endocrine profiling of reproductive status and evidence of pseudopregnancy in the Pacific walrus (Odobenus rosmarus divergens).

Endocrine profiling is an increasingly utilized tool for detecting pregnancies in wild populations of mammals. Given the difficulty in calculating reproductive rates of Pacific walruses (Odobenus rosmarus divergens) the use of endocrine techniques for determining pregnancy rates could be particularly useful for management of the population. The goals of this study were to 1) determine if progesterone and total estrogen concentrations in ovarian tissues of female walruses could be used to determine reproductive state and 2) determine if walruses undergo a functional postpartum estrus, as is seen in other pinnipeds. Ovaries were collected from female walruses (n = 13) hunted in subsistence hunts by Alaska Native communities. Females were categorized as postpartum, full-term pregnant, pregnant diapause or unbred. Total estrogen concentrations were greatest in unbred (n = 2) and pregnant (n = 2) females. Progesterone concentrations were also nominally larger in unbred (n = 2) than pregnant (n = 2) and postpartum (n = 9) animals. Small samples sizes precluded the use of statistical comparisons among groups. Corpora lutea tissue samples in this study did not reflect the presence of a postpartum estrus in the month of May as postpartum females yielded lower total estrogen concentrations than unbred or pregnant animals. Both unbred animals were in a state of pseudopregnancy, which has not been physiologically described for this species before. The progesterone profiles in late (59 ng/g) and early (140 ng/g) pregnancy were lower than expected and fell within the range of the postpartum females (36-210 ng/g), suggesting low production of the hormone by the corpus luteum during these phases of pregnancy. Profiling reproductive hormones in free-ranging walruses demonstrates that an endocrine approach may be a valuable tool for determining reproductive status of females, however increased sample sizes and time of year must be considered to accurately separate pregnant versus pseudopregnant individuals.

Pinnipeds orient and control their whiskers: a study on Pacific walrus, California sea lion and Harbor seal.

Whisker touch is an active sensory system. Previous studies in Pinnipeds have adopted relatively stationary tasks to judge tactile sensitivity, which may not accurately promote natural whisker movements and behaviours. This study developed a novel feeding task, termed fish sweeping to encourage whisker movements. Head and whisker movements were tracked from video footage in Harbor seal (Phoca vitulina), California sea lion (Zalophus californianus) and Pacific walrus (Odobenus rosmarus divergens). All species oriented their head towards the moving fish target and moved their whiskers during the task. Some species also engaged in whisker control behaviours, including head-turning asymmetry in the Pacific walrus, and contact-induced asymmetry in the Pacific walrus and California sea lion: behaviours that have only previously been observed in terrestrial mammals. This study confirms that Pinnipeds should be thought of as whisker specialists, and that whisker control (movement and positioning) is an important aspect of touch sensing in these animals, especially in sea lions and walruses. That the California sea lion controls whisker movement in relation to an object, and also had large values of whisker amplitude, spread and asymmetry, suggests that California sea lions are a promising model with which to further explore active touch sensing.

Facing each other: mammal mothers and infants prefer the position favouring right hemisphere processing.

The right hemisphere plays a crucial role in social processing. Human mothers show a robust left cradling/holding bias providing greater right-hemispheric involvement in the exchange of social information between mother and infant. Here, we demonstrate that a similar bias is evident in face-to-face spatial interactions in marine and terrestrial non-primate mammals. Walruses and Indian flying foxes showed a significant population-level preference for the position which facilitates the use of the left visual field in both mother and infant. This behavioural lateralization may have emerged owing to benefits conferred by the enhanced right-hemispheric social processing providing the mother and infant an optimal perception of each other.

SHORT DURATION IMMOBILIZATION OF ATLANTIC WALRUS ( ODOBENUS ROSMARUS ROSMARUS) WITH ETORPHINE, AND REVERSAL WITH NALTREXONE.

Forty adult, male Atlantic walruses ( Odobenus rosmarus rosmarus) were successfully immobilized for the attachment of global positioning system loggers on their tusks and collection of various biological samples. A standard dose of 7.8 mg etorphine was used for each animal, regardless of body size. All animals were reversed with an iv or im injection of 250 mg naltrexone, immediately after tag attachment. Twenty-seven of the animals were intubated and ventilated with 100% oxygen during the recovery period. The induction time was, on average, 4 min 51 sec ± 1 min 46 sec. Several animals had venous pH, and Pco2 levels that indicated severe acidosis and hypercarbia. All animals recovered within an average of 5 min 16 sec ± 2 min 47 sec after reversal. The total time from darting to recovery was 15 min 23 sec ± 3 min 33 sec. The use of naltrexone is recommended for reversal of etorphine immobilization in adult, male walruses, and the use of positive-pressure ventilation with oxygen is highly encouraged.

A comparison of blood nitric oxide metabolites and hemoglobin functional properties among diving mammals.

The ability of marine mammals to hunt prey at depth is known to rely on enhanced oxygen stores and on selective distribution of blood flow, but the molecular mechanisms regulating blood flow and oxygen transport remain unresolved. To investigate the molecular mechanisms that may be important in regulating blood flow, we measured concentration of nitrite and S-nitrosothiols (SNO), two metabolites of the vasodilator nitric oxide (NO), in the blood of 5 species of marine mammals differing in their dive duration: bottlenose dolphin, South American sea lion, harbor seal, walrus and beluga whale. We also examined oxygen affinity, sensitivity to 2,3-diphosphoglycerate (DPG) and nitrite reductase activity of the hemoglobin (Hb) to search for possible adaptive variations in these functional properties. We found levels of plasma and red blood cells nitrite similar to those reported for terrestrial mammals, but unusually high concentrations of red blood cell SNO in bottlenose dolphin, walrus and beluga whale, suggesting enhanced SNO-dependent signaling in these species. Purified Hbs showed similar functional properties in terms of oxygen affinity and sensitivity to DPG, indicating that reported large variations in blood oxygen affinity among diving mammals likely derive from phenotypic variations in red blood cell DPG levels. The nitrite reductase activities of the Hbs were overall slightly higher than that of human Hb, with the Hb of beluga whale, capable of longest dives, having the highest activity. Taken together, these results underscore adaptive variations in circulatory NO metabolism in diving mammals but not in the oxygenation properties of the Hb.

Decadal Bering Sea seascape change: consequences for Pacific walruses and indigenous hunters.

The most significant factors currently affecting the Pacific walrus (Odobenus rosmarus divergens) population are climate change and consequent changes in sea-ice morphology and dynamics. This paper integrates recent physical sea-ice change in the Bering Sea with biological and ecological conditions of walruses in their winter-spring reproductive habitat. Historically, walrus in winter-spring depended on a critical mass of sea-ice habitat to optimize social networking, reproductive fitness, feeding behavior, migration, and energetic efficiency. During 2003-2013, our cross-disciplinary, multiscale analysis from shipboard observations, satellite imagery, and ice-floe tracking, reinforced by information from indigenous subsistence hunters, documented change of sea-ice structure from a plastic continuum to a "mixing bowl" of ice floes moving more independently. This fragmentation of winter habitat preconditions the walrus population toward dispersal mortality and will also negatively affect the availability of resources for indigenous communities. We urge an expanded research and management agenda that integrates walrus natural history and habitat more completely with changing sea-ice morphology and dynamics at multiple scales, while also meeting the needs of local communities.

Zinc Isotope Ratios as Indicators of Diet and Trophic Level in Arctic Marine Mammals.

Carbon and nitrogen stable isotope ratios of bone collagen are an established method for dietary reconstruction, but this method is limited by the protein preservation. Zinc (Zn) is found in bioapatite and the isotopic compositions of this element constitute a very promising dietary indicator. The extent of fractionation of Zn isotopes in marine environments, however, remains unknown. We report here on the measurement of zinc, carbon and nitrogen isotopes in 47 marine mammals from the archaeological site of Arvik in the Canadian Arctic. We undertook this study to test and demonstrate the utility of Zn isotopes in recent mammal bone minerals as a dietary indicator by comparing them to other isotopic dietary tracers. We found a correlation between δ66Zn values and trophic level for most species, with the exception of walruses, which may be caused by their large seasonal movements. δ6Zn values can therefore be used as a dietary indicator in marine ecosystems for both modern and recent mammals.

Albicetus oxymycterus, a New Generic Name and Redescription of a Basal Physeteroid (Mammalia, Cetacea) from the Miocene of California, and the Evolution of Body Size in Sperm Whales.

Living sperm whales are represented by only three species (Physeter macrocephalus, Kogia breviceps and Kogia sima), but their fossil record provides evidence of an ecologically diverse array of different forms, including morphologies and body sizes without analog among living physeteroids. Here we provide a redescription of Ontocetus oxymycterus, a large but incomplete fossil sperm whale specimen from the middle Miocene Monterey Formation of California, described by Remington Kellogg in 1925. The type specimen consists of a partial rostrum, both mandibles, an isolated upper rostrum fragment, and incomplete tooth fragments. Although incomplete, these remains exhibit characteristics that, when combined, set it apart morphologically from all other known physeteroids (e.g., a closed mesorostral groove, and the retention of enameled tooth crowns). Kellogg originally placed this species in the genus Ontocetus, a enigmatic tooth taxon reported from the 19th century, based on similarities between the type specimen Ontocetus emmonsi and the conspicuously large lower dentition of Ontocetus oxymycterus. However, the type of the genus Ontocetus is now known to represent a walrus tusk (belonging to fossil Odobenidae) instead of a cetacean tooth. Thus, we assign this species to the new genus Albicetus, creating the new combination of Albicetus oxymycterus, gen. nov. We provide new morphological observations of the type specimen, including a 3D model. We also calculate a total length of approximately 6 m in life, using cranial proxies of body size for physeteroids. Lastly, a phylogenetic analysis of Albicetus oxymycterus with other fossil and living Physeteroidea resolves its position as a stem physeteroid, implying that large body size and robust dentition in physeteroids evolved multiple times and in distantly related lineages.

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). RESULTS: The 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. CONCLUSIONS: The best thermographic and most stable reference points for monitoring body temperature in marine mammals are open blowhole in cetaceans and eyes in pinnipeds.

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 13,808 ng/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 90,263 ng/g lipids, respectively. The specifics of diet as a source of pesticide accumulation in these two marine mammal species are discussed.

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.

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.

Underwater passive acoustic localization of Pacific walruses in the northeastern Chukchi Sea.

This paper develops and applies a linearized Bayesian localization algorithm based on acoustic arrival times of marine mammal vocalizations at spatially-separated receivers which provides three-dimensional (3D) location estimates with rigorous uncertainty analysis. To properly account for uncertainty in receiver parameters (3D hydrophone locations and synchronization times) and environmental parameters (water depth and sound-speed correction), these quantities are treated as unknowns constrained by prior estimates and prior uncertainties. Unknown scaling factors on both the prior and arrival-time uncertainties are estimated by minimizing Akaike's Bayesian information criterion (a maximum entropy condition). Maximum a posteriori estimates for sound source locations and times, receiver parameters, and environmental parameters are calculated simultaneously using measurements of arrival times for direct and interface-reflected acoustic paths. Posterior uncertainties for all unknowns incorporate both arrival time and prior uncertainties. Monte Carlo simulation results demonstrate that, for the cases considered here, linearization errors are small and the lack of an accurate sound-speed profile does not cause significant biases in the estimated locations. A sequence of Pacific walrus vocalizations, recorded in the Chukchi Sea northwest of Alaska, is localized using this technique, yielding a track estimate and uncertainties with an estimated speed comparable to normal walrus swim speeds.

Study of sleep in a walrus.

Several behavioral and physiological adaptations have been developed in evolution of Pinnipeds allowing them to sleep both on land and in water. To date sleep has been examined in detail in eared and true seals (the families of Otariidae and Phocidae). The aim of this study was to examine sleep in another semiaquatic mammal - the walrus, which is the only extant representative of the family Odobenidae. Slow wave and paradoxical sleep (SWS and PS) in the examined walrus (2 year old female, weight 130 kg) averaged 19.4 ± 2.0 and 6.9 ± 1.1% of 24-h when on land, and 20.5 ± 0.8% of 24-h and 1.1 ± 0.6% when in water, respectively. The average duration of PS episode was 6.4 ± 0.6 min (maximum 23 min) when on land and 1.8 ± 0.1 min (maximum 3.3 min) when in water. In water, sleep occurred predominantly while the walrus submerged and lay on the bottom of the pool (89% of total sleep time). The walrus usually woke up while emerging to the surface for breathing. Most often EEG slow waves developed synchronously in both cortical hemispheres (90% of SWS time when on land and 97% when in water). Short episodes of interhemispheric EEG asymmetry usually coincided with brief opening of one eye. The pattern of sleep in the walrus was similar to the pattern of sleep in the Otariidae seals while on land (predominantly bilateral SWS, accompanied by regular breathing) and to the pattern of sleep in the Phocidae while in water (sleep during apneas both in depth and at the surface, interrupted by brief arousal when emerging for breathing).

Tracking of Pacific walruses in the Chukchi Sea using a single hydrophone.

The vocal repertoire of Pacific walruses includes underwater sound pulses referred to as knocks and bell-like calls. An extended acoustic monitoring program was performed in summer 2007 over a large region of the eastern Chukchi Sea using autonomous seabed-mounted acoustic recorders. Walrus knocks were identified in many of the recordings and most of these sounds included multiple bottom and surface reflected signals. This paper investigates the use of a localization technique based on relative multipath arrival times (RMATs) for potential behavior studies. First, knocks are detected using a semi-automated kurtosis-based algorithm. Then RMATs are matched to values predicted by a ray-tracing model. Walrus tracks with vertical and horizontal movements were obtained. The tracks included repeated dives between 4.0 m and 15.5 m depth and a deep dive to the sea bottom (53 m). Depths at which bell-like sounds are produced, average knock production rate and source levels estimates of the knocks were determined. Bell sounds were produced at all depths throughout the dives. Average knock production rates varied from 59 to 75 knocks/min. Average source level of the knocks was estimated to 177.6 ± 7.5 dB re 1 µPa peak @ 1 m.