Subsurface swimming and stationary diving are metabolically cheap in adult Pacific walruses (Odobenus rosmarus divergens)

Walruses rely on sea-ice to efficiently forage and rest between diving bouts while maintaining proximity to prime foraging habitat. Recent declines in summer sea ice have resulted in walruses hauling out on land where they have to travel farther to access productive benthic habitat while potentially increasing energetic costs. Despite the need to better understand the impact of sea ice loss on energy expenditure, knowledge about metabolic demands of specific behaviours in walruses is scarce. In the present study, 3 adult female Pacific walruses (Odobenus rosmarus divergens) participated in flow-through respirometry trials to measure metabolic rates while floating inactive at the water surface during a minimum of 5 min, during a 180-second stationary dive, and while swimming horizontally underwater for ∼90 m. Metabolic rates during stationary dives (3.82±0.56 l O2 min−1) were lower than those measured at the water surface (4.64±1.04 l O2 min−1), which did not differ from rates measured during subsurface swimming (4.91±0.77 l O2 min−1). Thus, neither stationary diving nor subsurface swimming resulted in metabolic rates above those exhibited by walruses at the water surface. These results suggest that walruses minimize their energetic investment during underwater behaviours as reported for other marine mammals. Although environmental factors experienced by free-ranging walruses (e.g., winds or currents) likely affect metabolic rates, our results provide important information for understanding how behavioural changes affect energetic costs and can be used to improve bioenergetics models aimed at predicting the metabolic consequences of climate change on walruses.

Evaluation of Satellite Imagery for Monitoring Pacific Walruses at a Large Coastal Haulout

Pacific walruses (Odobenus rosmarus divergens) are using coastal haulouts in the Chukchi Sea more often and in larger numbers to rest between foraging bouts in late summer and autumn in recent years, because climate warming has reduced availability of sea ice that historically had provided resting platforms near their preferred benthic feeding grounds. With greater numbers of walruses hauling out in large aggregations, new opportunities are presented for monitoring the population. Here we evaluate different types of satellite imagery for detecting and delineating the peripheries of walrus aggregations at a commonly used haulout near Point Lay, Alaska, in 2018–2020. We evaluated optical and radar imagery ranging in pixel resolutions from 40 m to ~1 m: specifically, optical imagery from Landsat, Sentinel-2, Planet Labs, and DigitalGlobe, and synthetic aperture radar (SAR) imagery from Sentinel-1 and TerraSAR-X. Three observers independently examined satellite images to detect walrus aggregations and digitized their peripheries using visual interpretation. We compared interpretations between observers and to high-resolution (~2 cm) ortho-corrected imagery collected by a small unoccupied aerial system (UAS). Roughly two-thirds of the time, clouds precluded clear optical views of the study area from satellite. SAR was unaffected by clouds (and darkness) and provided unambiguous signatures of walrus aggregations at the Point Lay haulout. Among imagery types with 4–10 m resolution, observers unanimously agreed on all detections of walruses, and attained an average 65% overlap (sd 12.0, n 100) in their delineations of aggregation boundaries. For imagery with ~1 m resolution, overlap agreement was higher (mean 85%, sd 3.0, n 11). We found that optical satellite sensors with moderate resolution and high revisitation rates, such as PlanetScope and Sentinel-2, demonstrated robust and repeatable qualities for monitoring walrus haulouts, but temporal gaps between observations due to clouds were common. SAR imagery also demonstrated robust capabilities for monitoring the Point Lay haulout, but more research is needed to evaluate SAR at haulouts with more complex local terrain and beach substrates.

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.

Fecal Methylmercury Correlates With Gut Microbiota Taxa in Pacific Walruses (Odobenus rosmarus divergens)

ObjectivesMethylmercury metabolism was investigated in Pacific walruses (Odobenus rosmarus divergens) from St. Lawrence Island, Alaska, United States.MethodsTotal mercury and methylmercury concentrations were measured in fecal samples and paired colon samples (n = 16 walruses). Gut microbiota composition and diversity were determined using 16S rRNA gene sequencing. Associations between fecal and colon mercury and the 24 most prevalent gut microbiota taxa were investigated using linear models.ResultsIn fecal samples, the median values for total mercury, methylmercury, and %methylmercury (of total mercury) were 200 ng/g, 4.7 ng/g, and 2.5%, respectively, while in colon samples, the median values for the same parameters were 28 ng/g, 7.8 ng/g, and 26%, respectively. In fecal samples, methylmercury was negatively correlated with one Bacteroides genus, while members of the Oscillospirales order were positively correlated with both methylmercury and %methylmercury (of total mercury). In colon samples, %methylmercury (of total mercury) was negatively correlated with members of two genera, Romboutsia and Paeniclostridium.ConclusionsMedian %methylmercury (of total mercury) was 10 times higher in the colon compared to the fecal samples, suggesting that methylmercury was able to pass through the colon into systemic circulation. Fecal total mercury and/or methylmercury concentrations in walruses were comparable to some human studies despite differences in seafood consumption rates, suggesting that walruses excreted less mercury. There are no members (at this time) of the Oscillospirales order which are known to contain the genes to methylate mercury, suggesting the source of methylmercury in the gut was from diet and not in vivo methylation.

Lung function assessment in the Pacific walrus (Odobenus rosmarus divergens) while resting on land and submerged in water

In the present study, we examined lung function in healthy resting adult (born in 2003) Pacific walruses (Odobenus rosmarus divergens) by measuring respiratory flow (V̇) using a custom-made pneumotachometer. Three female walruses (670 – 1025 kg) voluntarily participated in spirometry trials while spontaneously breathing on land (sitting and lying down in sternal recumbency) and floating in water. While sitting, two walruses performed active respiratory efforts, and one animal participated in lung compliance measurements. For spontaneous breaths, V̇ was lower when lying down (e.g. expiration: 7.1±1.2 l · s−1) as compared to when in water (9.9±1.4 l · s−1), while tidal volume (VT, 11.5±4.6 l), breath duration (4.6±1.4 s), and respiratory frequency (7.6±2.2 breaths · min−1) remained the same. The measured VT and specific dynamic lung compliance (0.32±0.07 cmH2O−1) for spontaneous breaths, were higher than those estimated for similarly sized terrestrial mammals. The VT increased with body mass (allometric mass-exponent=1.29) and ranged from 3 to 43% of the estimated total lung capacity (TLCest) for spontaneous breaths. When normalized for TLCest, the maximal expiratory V̇ (V̇exp) was higher than that estimated in phocids, but lower than that reported in cetaceans and the California sea lion. The V̇exp was maintained over all lung volumes during spontaneous and active respiratory manoeuvres. We conclude that location (water or land) affects lung function in the walrus and should be considered when studying respiratory physiology in semi-aquatic marine mammals.

Determining sex of adult Pacific walruses from mandible measurements

Pacific walruses (Odobenus rosmarus divergens) play a vital role in Arctic marine ecosystems and the subsistence lifestyle of Alaska Native communities. Museum collections contain numerous archaeological and historic walrus specimens that have proven useful in a variety of studies; however, for many cases, the sex of these specimens is unknown. Sexes of adult (&gt; 5 years determined by tooth aging) Atlantic walruses (Odobenus rosmarus rosmarus) have been accurately determined in previous studies using mandible measurements. We tested the validity of this approach for Pacific walruses, and used full fusion of the mandibular symphysis to define adults. Using high precision digital calipers (± 0.01 mm), four measurements were taken either on the left or right side of 91 walrus mandibles: 80 modern mandibles (70 known-sex specimens; 10 unknown-sex specimens) and 11 archaeological mandibles of unknown sex. We used linear discriminant function analysis (LDFA) to determine what measurements best distinguished Pacific walrus males from females. Minimum mandible thickness had the most predictive power, whereas mandible length, height, and depth, were less predictive. Posterior probabilities indicated that LDFA classified the known-sex Pacific walruses with 100% accuracy, and unknown sex with ≥ 90% probability. The ability to define the sex of unknown individuals accurately could greatly increase the sample size of future projects dealing with skeletal remains, and will improve future research efforts.