Distribution and demographics of mysids (Crustacea: Mysida) as prey for gray whales (Eschrichtius robustus) in northwest Washington state.

Background: The movement and distribution of gray whales (Eschrichtius robustus) during the feeding season is likely dependent on the quality of foraging opportunities and the distribution of prey species. These dynamics are especially important to understand for the Pacific Coast Feeding Group (PCFG) of gray whales which spend the feeding season along the coast from northern California, USA through northern British Columbia, Canada. In Washington state, no previous work has been done to describe available gray whale prey. The main goal of this research was to initiate studies on an important gray whale prey item in northwest Washington, mysid shrimp (Mysida), by establishing a baseline understanding of mysid swarm demographics in the area and investigating patterns in gray whale and mysid presence. Methods: Prey samples were collected during June through November 2019 and June through September 2020 using a vertically-towed plankton net at seven sites in the Strait of Juan de Fuca and seven sites in the Pacific Ocean in areas where gray whales were known to feed. Mysids collected in the samples were counted and the sex, length, species, maturity, and gravidity were documented. Patterns in gray whale and mysid co-occurrence were explored through data visualization. Results: Seven species of mysids were observed in the survey area. In 2019, the number of mysids per tow increased steadily through the season, the most abundant species of mysids were Holmesimysis sculpta and Neomysis rayii, and sampled mysids averaged 4.7 mm in length. In 2020, mysids were abundant in tow samples in June and July but were not abundant in the remaining months of the sampling season. The average length of mysids in 2020 was 13.3 mm, and a large portion were sexually mature males and brooded females identified as H. sculpta. Throughout the survey area, the majority of whale sightings occurred later in the season in 2019 and earlier in the season in 2020, coinciding with the trends of sampled mysids. Discussion: This study provides the first description of mysid swarm composition and temporal variation in northwest Washington. Tows were dominated by a similar assemblage of mysid species as what is observed in other areas of the PCFG range. The differences in sampled mysid assemblages between years, and the presence of whales in the survey area in times and at sites where samples with high mysid counts were collected, suggest evidence for interesting predator-prey dynamics that warrant further investigation.

Assessment of a non-invasive approach to pregnancy diagnosis in gray whales through drone-based photogrammetry and faecal hormone analysis.

Knowledge of baleen whales' reproductive physiology is limited and requires long-term individual-based studies and innovative tools. We used 6 years of individual-level data on the Pacific Coast Feeding Group gray whales to evaluate the utility of faecal progesterone immunoassays and drone-based photogrammetry for pregnancy diagnosis. We explored the variability in faecal progesterone metabolites and body morphology relative to observed reproductive status and estimated the pregnancy probability for mature females of unknown reproductive status using normal mixture models. Individual females had higher faecal progesterone concentrations when pregnant than when presumed non-pregnant. Yet, at the population level, high overlap and variability in progesterone metabolite concentrations occurred between pregnant and non-pregnant groups, limiting this metric for accurate pregnancy diagnosis in gray whales. Alternatively, body width at 50% of the total body length (W50) correctly discriminated pregnant from non-pregnant females at individual and population levels, with high accuracy. Application of the model using W50 metric to mature females of unknown pregnancy status identified eight additional pregnancies with high confidence. Our findings highlight the utility of drone-based photogrammetry to non-invasively diagnose pregnancy in this group of gray whales, and the potential for improved data on reproductive rates for population management of baleen whales generally.

Influence of Life-History Parameters on Persistent Organic Pollutant Concentrations in Blubber of Eastern North Pacific Gray Whales (Eschrichtius robustus).

Exposure to persistent organic pollutants (POPs) can significantly impact marine mammal health, reproduction, and fitness. This study addresses a significant 20-year gap in gray whale contaminant monitoring through analysis of POPs in 120 blubber biopsies. The scope of this substantial sample set is noteworthy in its range and diversity with collection between 2003 and 2017 along North America's west coast and across diverse sex, age, and reproductive parameters, including paired mothers and calves. Mean blubber concentrations of polychlorinated biphenyls (∑PCBs), dichlorodiphenyltrichloroethanes (∑DDTs), and chlordanes (∑CHLs) generally decreased since previous reports (1968-1999). This is the first report of polybrominated diphenyl ethers (PBDEs) and select hexachlorocyclohexanes (HCHs) in this species. Statistical modeling of the 19 most frequently detected compounds in this dataset revealed sex-, age-, and reproductive status-related patterns, predominantly attributed to maternal offloading. Mean POP concentrations differed significantly by sex in adults (17 compounds, up to 3-fold higher in males) but not in immatures (all 19 compounds). Mean POP concentrations were significantly greater in adults versus immatures in both males (17 compounds, up to 12-fold) and females (13 compounds, up to 3-fold). POP concentrations were detected with compound-specific patterns in nursing calves, confirming maternal offloading for the first time in this species.

Benthic studies adjacent to Sakhalin Island, Russia, 2015 II: energy content of the zoobenthos in western gray whale feeding grounds.

The waters adjacent to the northeastern coast of Sakhalin Island, Russia, are an important feeding ground for the endangered western gray whale. Data on the energy available to foraging whales from their prey resources is required for researchers interested in modeling the bioenergetics of whale foraging, but little energy content information is available for the benthic prey communities of gray whales in this region. In this study, we describe the energy density (ED), biomass, and total energy availability (ED × biomass) of benthic prey sampled from two gray whale foraging areas adjacent to Sakhalin Island: the nearshore and offshore feeding areas. ED varied almost seven-fold among benthic taxa, ranging from 1.11 to 7.62 kJ/g wet mass. Although there was considerable variation within most prey groups, amphipods had the highest mean ED of all of groups examined (5.58 ± 1.44 kJ/g wet mass). Small sample sizes precluded us from detecting any seasonal or spatial differences in mean ED within or among taxa; however, mean biomass in the offshore feeding area was, in some cases, an order of magnitude higher than mean estimates in the nearshore feeding area, resulting in higher mean total energy available to foraging gray whales offshore (958-3313 kJ/m2) compared to nearshore (223-495 kJ/m2). While the proportion of total energy accounted for by amphipods was variable, this prey group generally made up a higher proportion of the total energy available in the benthos of the offshore feeding area than in the benthos of the nearshore feeding area. Data presented here will be used to inform bioenergetics modeling of the vital rates of mature females in an effort to improve understanding of population growth limits for western gray whales.

Gray whale density during seismic surveys near their Sakhalin feeding ground.

Oil and gas development off northeastern Sakhalin Island, Russia, has exposed the western gray whale population on their summer-fall foraging grounds to a range of anthropogenic activities, such as pile driving, dredging, pipeline installation, and seismic surveys. In 2015, the number of seismic surveys within a feeding season surpassed the level of the number and duration of previous seismic survey activities known to have occurred close to the gray whales' feeding ground, with the potential to cause disturbance to their feeding activity. To examine the extent that gray whales were potentially avoiding areas when exposed to seismic and vessel sounds, shore-based teams monitored the abundance and distribution of gray whales from 13 stations that encompassed the known nearshore feeding area. Gray whale density was examined in relation to natural (spatial, temporal, and prey energy) and anthropogenic (cumulative sound exposure from vessel and seismic sounds) explanatory variables using Generalized Additive Models (GAM). Distance from shore, water depth, date, and northing explained a significant amount of variation in gray whale densities. Prey energy from crustaceans, specifically amphipods, isopods, and cumaceans also significantly influenced gray whale densities in the nearshore feeding area. Increasing cumulative exposure to vessel and seismic sounds resulted in both a short- and longer-term decline in gray whale density in an area. This study provides further insights about western gray whale responses to anthropogenic activity in proximity to and within the nearshore feeding area. As the frequency of seismic surveys and other non-oil and gas anthropogenic activity are expected to increase off Sakhalin Island, it is critical to continue to monitor and assess potential impacts on this endangered population of gray whales.

Western gray whale behavioral response to seismic surveys during their foraging season.

Gray whales utilizing their foraging grounds off northeastern Sakhalin Island, Russia, have been increasingly exposed to anthropogenic activities related to oil and gas development over the past two decades. In 2015, four seismic vessels, contracted by two operators, conducted surveys near and within the gray whale feeding grounds. Mitigation and monitoring plans were developed prior to the survey and implemented in the field, with real-time data transfers to assist the implementation of measures aimed at minimizing impacts of acoustic exposure. This study examined the behavioral response of gray whales relative to vessel proximities and sounds generated during seismic exploration. Five shore-based teams monitored gray whale behavior from 1 June to 30 September using theodolite tracking and focal follow methodologies. Behavioral data were combined with acoustic and benthic information from studies conducted during the same period. A total of 1270 tracks (mean duration = 0.9 h) and 401 focal follows (1.1 h) were collected with gray whales exposed to sounds ranging from 59 to 172 dB re 1 µPa2 SPL. Mixed models were used to examine 13 movement and 10 respiration response variables relative to "natural," acoustic, and non-acoustic explanatory variables. Water depth and behavioral state were the largest predictors of gray whale movement and respiration patterns. As vessels approached whales with increasing seismic/vessel sound exposure levels and decreasing distances, several gray whale movement and respiration response variables significantly changed (increasing speed, directionality, surface time, respiration intervals, etc.). Although the mitigation measures employed could have reduced larger/long-term responses and sensitization to the seismic activities, this study illustrates that mitigation measures did not eliminate behavioral responses, at least in the short-term, of feeding gray whales to the activities.

Naturally stressed? Glucocorticoid profiles in blubber of blue and gray whales in response to life history parameters.

The goal of the present study was to carry out a thorough methodological validation and describe baseline profiles for glucocorticoid hormones (cortisol and corticosterone) in blubber from blue (n = 77) and gray (n = 103) whales from the eastern North Pacific Ocean. For each species, we modelled cortisol and corticosterone concentrations in response to life history parameters (age, sex, reproductive status) and season or geographic location. In blue whales, cortisol concentrations did not vary significantly by age class, sex, or reproductive status, whereas corticosterone was significantly lower in immature than in adult females (p < .001). In gray whales, cortisol concentrations were significantly higher in lactating whales (p < .05), while corticosterone was significantly different between females and males (p = .001) and elevated in calves (p = .003). In gray whales, corticosterone concentrations were significantly lower in males sampled later in the year (August to November) compared to both sexes sampled between March and August (p = .05), but no seasonal trend occurred in blue whales. Our results indicate that glucocorticoid actions vary between species and sex in large whales. Analysis of multiple hormones improves our understanding of the physiology of maintaining metabolic homeostasis or coping with chronic stressors.

Assessment of fecal steroid and thyroid hormone metabolites in eastern North Pacific gray whales.

Baleen whale fecal samples have high potential for endocrine monitoring, which can be used as a non-invasive tool to identify the physiological response to disturbance events and describe population health and vital rates. In this study, we used commercial enzyme-linked immunosorbent assays to validate and quantify fecal steroid (progestins, androgens and glucocorticoids) and thyroid hormone metabolite concentrations in eastern North Pacific gray whales (Eschrichtius robustus) along the Oregon coast, USA, from May to October of 2016-2018. Higher mean progestin metabolite concentrations were observed in postweaning females, followed by pregnant females. Mean androgen, glucocorticoid and thyroid metabolites were higher in mature males. Progestin, glucocorticoids and thyroid fecal metabolites varied significantly by year, with positive correlations between progestin and androgen, and between glucocorticoid and thyroid metabolites. We also present two case studies of a documented injured whale and a mature male displaying reproductive competitive behavior, which provide reference points for physiologically stressed individuals and adult breeding males, respectively. Our methods and findings advance the knowledge of baleen whale physiology, can help guide future research on whale physiology and can inform population management and conservation efforts regarding minimizing the impact of anthropogenic stressors on whales.

Growth and development of North Pacific gray whales (Eschrichtius robustus).

Understanding variability in growth patterns of marine mammals provides insights into the health of individuals and status of populations. Body growth of gray whales (Eschrichtius robustus) has been described for particular life stages, but has not been quantified across all ages. We derived a comprehensive growth equation for gray whales by fitting a two-phased growth model to age-specific length data of eastern North Pacific gray whales that were captured, stranded, or harvested between 1926 and 1997. To predict mass-at-age, we used the allometric relationship between mass and length. We found that on average (± SD), calves were 4.6 ± 0.043 m and 972 ± 26 kg at birth, and reached 8.5 ± 0.095 m and 6,019 ± 196 kg by the end of their first year of life (n = 118). Thus, calves almost double (2×) in length and octuple (8×) in mass while nursing, and are effectively about two-thirds of their asymptotic adult length and one-third of their maximum mass when weaned. The large sample of aged individuals (n = 730) indicates that gray whales live up to ~48 years and have a life expectancy of < 30 years. Adult females attain a mean (± SD) asymptotic size of 13.1 ± 0.048 m and 20,758 ± 222 kg, while the smaller males average 12.6 ± 0.048 m and 19,938 ± 222 kg at ~40 years of age. Females are thereby ~4% longer and heavier than males. These age-specific estimates of body size can be used to estimate food requirements and assess nutritional status of individuals.

Gray whale transcriptome reveals longevity adaptations associated with DNA repair and ubiquitination.

One important question in aging research is how differences in genomics and transcriptomics determine the maximum lifespan in various species. Despite recent progress, much is still unclear on the topic, partly due to the lack of samples in nonmodel organisms and due to challenges in direct comparisons of transcriptomes from different species. The novel ranking-based method that we employ here is used to analyze gene expression in the gray whale and compare its de novo assembled transcriptome with that of other long- and short-lived mammals. Gray whales are among the top 1% longest-lived mammals. Despite the extreme environment, or maybe due to a remarkable adaptation to its habitat (intermittent hypoxia, Arctic water, and high pressure), gray whales reach at least the age of 77 years. In this work, we show that long-lived mammals share common gene expression patterns between themselves, including high expression of DNA maintenance and repair, ubiquitination, apoptosis, and immune responses. Additionally, the level of expression for gray whale orthologs of pro- and anti-longevity genes found in model organisms is in support of their alleged role and direction in lifespan determination. Remarkably, among highly expressed pro-longevity genes many are stress-related, reflecting an adaptation to extreme environmental conditions. The conducted analysis suggests that the gray whale potentially possesses high resistance to cancer and stress, at least in part ensuring its longevity. This new transcriptome assembly also provides important resources to support the efforts of maintaining the endangered population of gray whales.

Comparative Respiratory Physiology in Cetaceans.

In the current study, we used breath-by-breath respirometry to evaluate respiratory physiology under voluntary control in a male beluga calf [Delphinapterus leucas, body mass range (M b): 151-175 kg], an adult female (estimated M b = 500-550 kg) and a juvenile male (M b = 279 kg) false killer whale (Pseudorca crassidens) housed in managed care. Our results suggest that the measured breathing frequency (f R) is lower, while tidal volume (V T) is significantly greater as compared with allometric predictions from terrestrial mammals. Including previously published data from adult bottlenose dolphin (Tursiops truncatus) beluga, harbor porpoise (Phocoena phocoena), killer whale (Orcinus orca), pilot whale (Globicephala scammoni), and gray whale (Eschrichtius robustus) show that the allometric mass-exponents for V T and f R are similar to that for terrestrial mammals (V T: 1.00, f R: -0.20). In addition, our results suggest an allometric relationship for respiratory flow ( V . ), with a mass-exponent between 0.63 and 0.70, and where the expiratory V . was an average 30% higher as compared with inspiratory V . . These data provide enhanced understanding of the respiratory physiology of cetaceans and are useful to provide proxies of lung function to better understand lung health or physiological limitations.

Recent dating of extinct Atlantic gray whale fossils, (Eschrichtius robustus), Georgia Bight and Florida, western Atlantic Ocean.

The Atlantic gray whale (Eschrichtius robustus) presents an interesting case study of climate related dispersal and extinction. While (limited) fossil records confirm its presence in the Atlantic up until the 18th Century, its abundance and distribution within the Eastern and Western basins are still not well understood. The discovery of presumed gray whale fossil remains from the Georgia Bight and the Atlantic coast of Florida, from the mid-1980s to late-2000s, provides a new opportunity to recover additional data regarding their chronology within the Western basin. Here, we apply accelerator mass spectrometry radiocarbon techniques to six fossil whale finds, identifying dates within marine isotope stage 3 (59-24 ka) and the late Holocene, ∼2,000 yr BP. We additionally confirm the taxonomic identification of two fossil bone samples as E. robustus using collagen peptide mass fingerprinting (ZooMS). The obtained dates, when combined with a larger corpus of previously published Atlantic gray whale fossil dates, support the hypothesis for the decline of the Atlantic gray whale in the late Pleistocene and the late Holocene. These new data augment the findings of the Eastern Atlantic Basin and better incorporate the Western Atlantic Basin into a pan-ocean understanding for the species.

Ampelisca eschrichtii Krøyer, 1842 (Ampeliscidae) of the Sakhalin Shelf in the Okhotsk Sea starve in summer and feast in winter.

BACKGROUND: Ampelisca eschrichtii Krøyer, 1842 of the Sakhalin Shelf of the Okhotsk Sea, Far Eastern Russia, comprise the highest known biomass concentration of any amphipod population in the world and are a critically important prey source for western gray whales. Growth and reproduction in this population has not been apparent in summer. However, they are not accessible for sampling in winter to test a previous default conclusion that they grow and reproduce in winter. METHODS: We tested the default winter growth and reproduction hypothesis by detailed comparisons of the brood and gonad development among 40 females and 14 males and brood sizes among females observed since 2002. Our test included six predictions of reproductive synchrony that would be apparent from gonad and brood morphology if active reproduction occurs in summer. RESULTS: We found high prevalences of undersized and damaged oocytes, undersized broods, a lack of females brooding fully formed juveniles, atrophied ovaries, and males with mature sperm but lacking fully developed secondary sex morphologies required for pelagic mating. All of these conditions are consistent with trophic stress and starvation. DISCUSSION: These A. eschrichtii populations therefore appear to starve in summer and to grow and reproduce in winter. The Offshore A. eschrichtii populations occur in summer below water strata bearing high phytoplankton biomasses. These populations are more likely to feed successfully in winter when storms mix phytoplankton to their depths.

Characterization of the Gray Whale Eschrichtius robustus Genome and a Genotyping Array Based on Single-Nucleotide Polymorphisms in Candidate Genes.

Genetic and genomic approaches have much to offer in terms of ecology, evolution, and conservation. To better understand the biology of the gray whale Eschrichtius robustus (Lilljeborg, 1861), we sequenced the genome and produced an assembly that contains ∼95% of the genes known to be highly conserved among eukaryotes. From this assembly, we annotated 22,711 genes and identified 2,057,254 single-nucleotide polymorphisms (SNPs). Using this assembly, we generated a curated list of candidate genes potentially subject to strong natural selection, including genes associated with osmoregulation, oxygen binding and delivery, and other aspects of marine life. From these candidate genes, we queried 92 autosomal protein-coding markers with a panel of 96 SNPs that also included 2 sexing and 2 mitochondrial markers. Genotyping error rates, calculated across loci and across 69 intentional replicate samples, were low (0.021%), and observed heterozygosity was 0.33 averaged over all autosomal markers. This level of variability provides substantial discriminatory power across loci (mean probability of identity of 1.6 × 10-25 and mean probability of exclusion >0.999 with neither parent known), indicating that these markers provide a powerful means to assess parentage and relatedness in gray whales. We found 29 unique multilocus genotypes represented among our 36 biopsies (indicating that we inadvertently sampled 7 whales twice). In total, we compiled an individual data set of 28 western gray whales (WGSs) and 1 presumptive eastern gray whale (EGW). The lone EGW we sampled was no more or less related to the WGWs than expected by chance alone. The gray whale genomes reported here will enable comparative studies of natural selection in cetaceans, and the SNP markers should be highly informative for future studies of gray whale evolution, population structure, demography, and relatedness.

De novo assembling and primary analysis of genome and transcriptome of gray whale Eschrichtius robustus.

BACKGROUND: Gray whale, Eschrichtius robustus (E. robustus), is a single member of the family Eschrichtiidae, which is considered to be the most primitive in the class Cetacea. Gray whale is often described as a "living fossil". It is adapted to extreme marine conditions and has a high life expectancy (77 years). The assembly of a gray whale genome and transcriptome will allow to carry out further studies of whale evolution, longevity, and resistance to extreme environment. RESULTS: In this work, we report the first de novo assembly and primary analysis of the E. robustus genome and transcriptome based on kidney and liver samples. The presented draft genome assembly is complete by 55% in terms of a total genome length, but only by 24% in terms of the BUSCO complete gene groups, although 10,895 genes were identified. Transcriptome annotation and comparison with other whale species revealed robust expression of DNA repair and hypoxia-response genes, which is expected for whales. CONCLUSIONS: This preliminary study of the gray whale genome and transcriptome provides new data to better understand the whale evolution and the mechanisms of their adaptation to the hypoxic conditions.

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.

Critically endangered western gray whales migrate to the eastern North Pacific.

Western North Pacific gray whales (WGWs), once considered extinct, are critically endangered with unknown migratory routes and reproductive areas. We attached satellite-monitored tags to seven WGWs on their primary feeding ground off Sakhalin Island, Russia, three of which subsequently migrated to regions occupied by non-endangered eastern gray whales (EGWs). A female with the longest-lasting tag visited all three major EGW reproductive areas off Baja California, Mexico, before returning to Sakhalin Island the following spring. Her 22 511 km round-trip is the longest documented mammal migration and strongly suggests that some presumed WGWs are actually EGWs foraging in areas historically attributed to WGWs. The observed migration routes provide evidence of navigational skills across open water that break the near-shore north-south migratory paradigm of EGWs. Despite evidence of genetic differentiation, these tagging data indicate that the population identity of whales off Sakhalin Island needs further evaluation.

Introduction to the anatomy of the head of a neonate gray whale (Mysticeti, Eschrichtius robustus).

The gray whale (Eschrichtius robustus) is the sole living representative of the mysticete (baleen whale) family Eschrichtiidae. Previous anatomical work on gray whales has been limited owing, in part, to difficulties of specimen access. These contributions to the anatomy of the gray whale head based on dissection of a stranded specimen from northern California include detailed investigation of internal and external features that confirm existing information and provide new evidence for their functional roles, particularly in thermoregulation and feeding.

Passive restriction of blood flow and counter-current heat exchange via lingual retia in the tongue of a neonatal gray whale Eschrichtius robustus (Cetacea, Mysticeti).

Retia mirabilia play broad roles in cetacean physiology, including thermoregulation during feeding and pressure regulations during diving. Vascular bundles of lingual retia are described within the base of the tongue of a neonatal female gray whale (Eschrichtius robustus). Each rete consists of a central artery surrounded by four to six smaller veins. The retia and constituent vessels decrease in diameter as they extend anteriorly within the hyoglossus muscle from a position anterior to the basihyal cartilage toward the apex of the tongue. The position of the retia embedded in the hyoglossus and the anterior constriction of the vessels differs from reports of similar vascular bundles that were previously identified in gray whales. The retia likely serve as a counter-current heat exchange system to control body temperature during feeding. Cold blood flowing toward the body center within the periarterial veins would accept heat from warm blood in the central artery flowing toward the anterior end of the tongue. Although thermoregulatory systems have been identified within the mouths of a few mysticete species, the distribution of such vascular structures likely is more widespread among baleen whales than has previously been described.

Eye, nose, hair, and throat: external anatomy of the head of a neonate gray whale (Cetacea, Mysticeti, Eschrichtiidae).

Information is scarce on gray whale (Eschrichtius robustus) anatomy and that of mysticetes in general. Dissection of the head of a neonatal gray whale revealed novel anatomical details of the eye, blowhole, incisive papilla with associated nasopalatine ducts, sensory hairs, and throat grooves. Compared to a similar sized right whale calf, the gray whale eyeball is nearly twice as long. The nasal cartilages of the gray whale, located between the blowholes, differ from the bowhead in having accessory cartilages. A small, fleshy incisive papilla bordered by two blind nasopalatine pits near the palate's rostral tip, previously undescribed in gray whales, may be associated with the vomeronasal organ, although histological evidence is needed for definitive identification. Less well known among mysticetes are the numerous elongated, stiff sensory hairs (vibrissae) observed on the gray whale rostrum from the ventral tip to the blowhole and on the mandible. These hairs are concentrated on the chin, and those on the lower jaw are arranged in a V-shaped pattern. We confirm the presence of two primary, anteriorly converging throat grooves, confined to the throat region similar to those of ziphiid and physeteroid odontocetes. A third, shorter groove occurs lateral to the left primary groove. The throat grooves in the gray whale have been implicated in gular expansion during suction feeding.