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.

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.

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.

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.

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.

Enhancing the accuracy and efficiency of Pacific walrus (Odobenus rosmarus divergens) surveys: A comparison of visual and aerial imagery-based counts at coastal haulouts.

This study presents a semi-automated approach utilizing unoccupied aerial vehicle (UAV) surveys to accurately estimate the abundance of Pacific walruses at large coastal haulouts in Chukotka, Russia. Seven major haulout sites were surveyed during the summers and falls of 2017-2019. Walrus counts were performed using three distinct methods: traditional visual land-based counts, complete head counts utilizing georeferenced UAV imagery, and counting walruses within model polygons within the haulout outline and employing various extrapolation techniques to predict walrus abundance across the haulout area. The results indicated that traditional visual counts neither yielded consistent results nor allowed for uncertainty estimation, unlike the site- and date-specific direct extrapolation method and the non-specific linear regression model. These latter methods consistently provided estimates, on average, within 5% of the "true" abundance determined through complete photo-based head counts. Beside yielding accurate estimates, these semi-automated methods significantly reduced counting time by at least 63%, in contrast to complete head counts. The non-specific model, which allowed the estimation of walrus abundance based on the type of the terrain and the haulout area was less accurate compared with site and date specific estimates, but provided a tool to estimate abundance when no field visits are conducted, e.g., by using high-resolution satellite imagery to measure haulout area. This model revealed that the haulouts located on flat sandy beaches exhibited mean walrus densities approximately 30.5% times higher than those on rocky shores surrounded by cliffs: 0.879 (SD = 0.1302) and 0.648 (SD = 0.1753) walrus per m2 correspondingly. The estimated daily walrus abundance at major Chukotkan haulouts in 2017-2019 ranged between 15 and 94,660 (mean = 10,397, SD = 14,477) walruses with the maximum seasonal abundances reported at Cape Serdtse-Kamen as 94,960 on 10-Oct-2017, 26,850 on 10-Oct-2018, and 87,595 on 10-Oct-2019.

Interannual site fidelity by Svalbard walruses.

The Arctic is experiencing rapid reductions in sea ice, affecting all ice-dependant species. In the present study we examine interannual seasonal movements and habitat use in relation to sea ice coverage for one of the Arctic endemic marine mammals. We tagged 40 male walruses (Odobenus rosmarus) in the Svalbard Archipelago with custom-designed tusk-mounted GPS loggers. Twelve of these animals provided tracks that lasted 1-6 years. Eleven of the walruses displayed clear seasonal migratory behaviour between summer foraging areas and winter breeding areas. Individuals showed high inter-individual variation, but clear site fidelity, using the same areas in consecutive years despite variable sea ice conditions. The walruses swam 5225-10,406 km per year and travelled remarkably similar distances between years on an individual basis. The phenology of migration was not impacted by sea ice concentrations or daylight length but was consistent at the individual level, suggesting endogenous drivers. Sea ice concentrations influenced movement behaviour with animals showing more tortuous paths when in areas with heavy sea ice, possibly searching for polynyas where females reside. Ongoing climate change is expected to drastically change walrus habitat, and it remains to be seen if walruses will be able to shift from their fixed seasonal migratory routines.

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.

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.

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.

Mother-calf vocal communication in Atlantic walrus: a first field experimental study.

In all colonial pinnipeds studied, mother-young vocal recognition exists and allows rapid and reliable meetings in spite of the confusing environment of the breeding colony. The efficiency of this recognition process guarantees pup survival, especially in species where females alternate foraging sea trips and lactation periods on land. The Atlantic Walrus (Odobenus rosmarus rosmarus) is a highly gregarious pinniped with females attending their calves for an extended period of time (2-3 years). Although we expect mother-calf vocal recognition to occur in this species due to the high density of individuals packed in herds, it has never been experimentally demonstrated. Here, we assessed the individual stereotypy of both mother and calf barks recorded in the wild by measuring frequency and temporal acoustic parameters. Both discriminant function and artificial neural network analyses resulted in high correct classification rates, underlying a well-defined individual stereotypy in parameters related to frequency modulation and frequency values. Playback experiments showed that mothers were more responsive to the barks of their own calf than to those of unrelated young. Finally, propagation experiments revealed that barks propagate at greater distances over water surface than over ice, acoustic features such as frequency modulation and frequency spectrum being highly resistant to degradation during propagation. Thus, acoustic analysis and propagation experiments suggest that these frequency parameters might be the key acoustic features involved in the individual identification process. This experimental study clearly demonstrates that Atlantic walrus has developed a highly reliable mother-calf vocal communication allowing such strong social bond.

The influence of feeding, enrichment, and seasonal context on the behavior of Pacific Walruses (Odobenus rosmarus divergens).

Though some research exists concerning general behavior and activity patterns of Walruses in zoos or aquariums, less is known about how these patterns change in response to various environmental and temporal contexts. This study presents two studies assessing behavioral changes in relation to feeding period, object enrichment (OE), and season in a social group of four Pacific Walruses at the New York Aquarium. Study 1 examined behavior in relation to feeding context (nonfeed, prefeed, postfeed); data were collected over a three-week period, resulting in 47 observation sessions for each feeding context. Study 2 examined behavior in relation to OE and season; data were collected in two phases resulting in 12 enrichment and 9 no-enrichment (NE) observation sessions (Phase 1), and 21 enrichment and 18 NE observation sessions (Phase 2). Study 1 showed that after feeding, oral behavior increased while social behavior and total swim frequency decreased. In Study 2, both swim frequency and social behavior were found to interact with OE and phase, while oral behavior remained constant across all conditions. As in the wild, both studies found all animals to be swimming the majority of the time. Though every animal spent much of its swim time engaged in an Individual Swimming Pattern (ISP), both studies showed that the proportion of ISP (in relation to total time swimming) remained stable across all contexts, suggesting a potential functional role of the ISPs. These results are discussed in light of the ongoing debate over the role of stereotypies in welfare assessment.

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.

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.