Three decades of nearshore surveys reveal long-term patterns in gray whale habitat use, distribution, and abundance in the Northern California Current.

The nearshore waters of the Northern California Current support an important seasonal foraging ground for Pacific Coast Feeding Group (PCFG) gray whales. We examine gray whale distribution, habitat use, and abundance over 31 years (1992-2022) using standardized nearshore (< 5 km from shore) surveys spanning a large swath of the PCFG foraging range. Specifically, we generated density surface models, which incorporate detection probability into generalized additive models to assess environmental correlates of gray whale distribution and predict abundance over time. We illustrate the importance of coastal upwelling dynamics, whereby increased upwelling only yields higher gray whale density if interspersed with relaxation events, likely because this combination optimizes influx and retention of nutrients to support recruitment and aggregation of gray whale prey. Several habitat features influence gray whale distribution, including substrate, shelf width, prominent capes, and river estuaries. However, the influence of these features differs between regions, revealing heterogeneity in habitat preferences throughout the PCFG foraging range. Predicted gray whale abundance fluctuated throughout our study period, but without clear directional trends, unlike previous abundance estimates based on mark-recapture models. This study highlights the value of long-term monitoring, shedding light on the impacts of variable environmental conditions on an iconic nearshore marine predator.

The evolution of menopause in toothed whales.

Understanding how and why menopause has evolved is a long-standing challenge across disciplines. Females can typically maximize their reproductive success by reproducing for the whole of their adult life. In humans, however, women cease reproduction several decades before the end of their natural lifespan1,2. Although progress has been made in understanding the adaptive value of menopause in humans3,4, the generality of these findings remains unclear. Toothed whales are the only mammal taxon in which menopause has evolved several times5, providing a unique opportunity to test the theories of how and why menopause evolves in a comparative context. Here, we assemble and analyse a comparative database to test competing evolutionary hypotheses. We find that menopause evolved in toothed whales by females extending their lifespan without increasing their reproductive lifespan, as predicted by the 'live-long' hypotheses. We further show that menopause results in females increasing their opportunity for intergenerational help by increasing their lifespan overlap with their grandoffspring and offspring without increasing their reproductive overlap with their daughters. Our results provide an informative comparison for the evolution of human life history and demonstrate that the same pathway that led to menopause in humans can also explain the evolution of menopause in toothed whales.

The oldest mysticete in the Northern Hemisphere.

Extant baleen whales (Mysticeti) uniquely use keratinous baleen for filter-feeding and lack dentition, but the fossil record clearly shows that "toothed" baleen whales first appeared in the Late Eocene.1 Globally, only two Eocene mysticetes have been found, and both are from the Southern Hemisphere: Mystacodon selenensis from Peru, 36.4 mega-annum (Ma) ago1,2 and Llanocetus denticrenatus from Antarctica, 34.2 Ma ago.3,4 Based on a partial skull from the lower part of the Lincoln Creek Formation in Washington State, USA, we describe the Northern Hemisphere's geochronologically earliest mysticete, Fucaia humilis sp. nov. Geology, biostratigraphy, and magnetostratigraphy places Fucaia humilis sp. nov. in the latest Eocene (ca. 34.5 Ma ago, near the Eocene/Oligocene transition at 33.9 Ma ago), approximately coeval with the oldest record of fossil kelps, also in the northeastern Pacific.5 This observation leads to our hypothesis that the origin and development of a relatively stable, nutrient-rich kelp ecosystem5,6 in the latest Eocene may have fostered the radiation of small-sized toothed mysticetes (Family Aetiocetidae) in the North Pacific basin, a stark contrast to the larger Llanocetidae (whether Mystacodon belongs to llanocetids or another independent clade remains unresolved) with the latest Eocene onset of the Antarctic Circumpolar Current in the Southern Hemisphere.7,8,9 Our discovery suggests that disparate mechanisms and ecological scenarios may have nurtured contrasting early mysticete evolutionary histories in the Northern and Southern hemispheres.

Analysis of protected species observer data: Strengths, weaknesses, and application in the assessment of marine mammal responses to seismic surveys in the northern Gulf of Mexico 2002-2015.

Visual observation data collected by protected species observers (PSOs) is required per regulations stipulated in Notices to Lessees (NTLs) and geophysical survey Permits (Form BOEM-0328) issued to seismic operators in the Gulf of Mexico (GOM). Here, data collected by certified and trained PSOs during seismic surveys conducted between 2002-2015 were compiled and analyzed to assess utility in assessing marine mammal responses to seismic noise and effectiveness of required mitigation measures. A total of 3,886 agency-required bi-weekly PSO Effort and Sightings reports were analyzed comprising 598,319 hours of PSO visual effort and 15,117 visual sighting records of marine mammals. The observed closest point of approach (CPA) distance was statistically compared across five species groupings for four airgun activity levels (full, minimum source, ramp up, silent). Whale and dolphin detections were significantly farther from airgun array locations during full power operations versus silence, indicating some avoidance response to full-power operations. Dolphin CPA distances were also significantly farther from airguns operating at minimum source than silence. Blackfish were observed significantly farther from the airgun array during ramp up versus both full and minimum source activities. Blackfish were observed significantly closer to the airgun array during silent activities versus at full, minimum source, and ramp up activities. Beaked whales had the largest mean CPA for detection distance compared to all other species groups. Detection distances for beaked whales were not significantly differences between full and silent operations; however, the sample size was very low. Overall results are consistent with other studies indicating that marine mammals may avoid exposure to airgun sounds based on observed distance from the seismic source during specified source activities. There was geographic variability in sighting rates associated with specific areas of interest within the GOM. This study demonstrates that agency required PSO reports provide a robust and useful data set applicable to impact assessments; management, policy and regulatory decision making; and qualitative input for regional scientific, stock assessment and abundance studies. However, several improvements in content and consistency would facilitate finer-scale analysis of some topics (e.g., effort associated with specific activities, observer biases, sound field estimation) and support statistical comparisons that could provide further insight into marine mammal responses and mitigation efficacy.

Evolutionary novelties underlie sound production in baleen whales.

Baleen whales (mysticetes) use vocalizations to mediate their complex social and reproductive behaviours in vast, opaque marine environments1. Adapting to an obligate aquatic lifestyle demanded fundamental physiological changes to efficiently produce sound, including laryngeal specializations2-4. Whereas toothed whales (odontocetes) evolved a nasal vocal organ5, mysticetes have been thought to use the larynx for sound production1,6-8. However, there has been no direct demonstration that the mysticete larynx can phonate, or if it does, how it produces the great diversity of mysticete sounds9. Here we combine experiments on the excised larynx of three mysticete species with detailed anatomy and computational models to show that mysticetes evolved unique laryngeal structures for sound production. These structures allow some of the largest animals that ever lived to efficiently produce frequency-modulated, low-frequency calls. Furthermore, we show that this phonation mechanism is likely to be ancestral to all mysticetes and shares its fundamental physical basis with most terrestrial mammals, including humans10, birds11, and their closest relatives, odontocetes5. However, these laryngeal structures set insurmountable physiological limits to the frequency range and depth of their vocalizations, preventing them from escaping anthropogenic vessel noise12,13 and communicating at great depths14, thereby greatly reducing their active communication range.

A tradeoff evolution between acoustic fat bodies and skull muscles in toothed whales.

Toothed whales have developed specialized echolocation abilities that are crucial for underwater activities. Acoustic fat bodies, including the melon, extramandibular fat body, and intramandibular fat body, are vital for echolocation. This study explores the transcriptome of acoustic fat bodies in toothed whales, revealing some insight into their evolutionary origins and ecological significance. Comparative transcriptome analysis of acoustic fat bodies and related tissues in a harbor porpoise and a Pacific white-sided dolphin reveals that acoustic fat bodies possess characteristics of both muscle and adipose tissue, occupying an intermediate position. The melon and extramandibular fat body exhibit specific muscle-related functions, implying an evolutionary connection between acoustic fat bodies and muscle tissue. Furthermore, we suggested that the melon and extramandibular fat body originate from intramuscular adipose tissue, a component of white adipose tissue. The extramandibular fat body has been identified as an evolutionary homolog of the masseter muscle, supported by the specific expression of MYH16, a pivotal protein in masticatory muscles. The intramandibular fat body, located within the mandibular foramen, shows possibilities of the presence of several immune-related functions, likely due to its proximity to bone marrow. Furthermore, this study sheds light on leucine modification in the catabolic pathway, which leads to the accumulation of isovaleric acid in acoustic fat bodies. Swallowing without chewing, a major toothed whale feeding ecology adaptation, makes the masticatory muscle redundant and leads to the formation of the extramandibular fat body. We propose that the intramuscular fat enlargement in facial muscles, which influences acoustic fat body development, is potentially related to the substantial reorganization of head morphology in toothed whales during aquatic adaptation.

Melon and rostral muscle morphology of Gervais' beaked whale (Mesoplodon europaeus): Alternating patterns of bilateral asymmetry.

Toothed whales utilize specialized nasal structures such as the lipid-rich melon to produce sound and propagate it into the aquatic environment. Very little nasal morphology of mesoplodont beaked whales has been described in the literature, and the anatomy of the melon and associated musculature of Gervais' beaked whale (Mesoplodon europaeus) remains undescribed. Heads of three (n = 3) Gervais' beaked whales were examined in detail via dissection as well as computed tomography (CT) and magnetic resonance imaging (MRI). Two additional Gervais' beaked whale individuals (n = 2) were studied via archived CT and MRI scans. Representative transverse dissection sections of the melon were processed for polarized light imaging to verify the presence of tendons inserting into the melon tissue. Three-dimensional (3D) CT reconstructions of the melon, rostral muscles, and associated structures were performed to assess morphology and spatial relationships. In all individuals, the melon's main body demonstrated a bilaterally asymmetrical, curvilinear geometry. This curvilinear shape was defined by a pattern of alternating asymmetry in the medial rostral muscles that projected into the melon's tissue. In transverse polarized light imaging, a network of tendons originating from these asymmetrical rostral muscle projections was observed permeating the melon's lipid tissue. This curvilinear melon morphology and associated asymmetrical musculature suggest a means of lengthening the lipid pathway within a relatively short dimensional footprint. In addition, the species-specific arrangement of muscular projections suggests complex fine-tuning of the melon's geometry during echolocation. Further studies may lend additional insight into the function of this unusual melon morphology.

Convergence in hearing-related genes between echolocating birds and mammals.

Echolocation, the detection of objects by means of sound waves, has evolved independently in diverse animals. Echolocators include not only mammals such as toothed whales and yangochiropteran and rhinolophoid bats but also Rousettus fruit bats, as well as two bird lineages, oilbirds and swiftlets. In whales and yangochiropteran and rhinolophoid bats, positive selection and molecular convergence has been documented in key hearing-related genes, such as prestin (SLC26A5), but few studies have examined these loci in other echolocators. Here, we examine patterns of selection and convergence in echolocation-related genes in echolocating birds and Rousettus bats. Fewer of these loci were under selection in Rousettus or birds compared with classically recognized echolocators, and elevated convergence (compared to outgroups) was not evident across this gene set. In certain genes, however, we detected convergent substitutions with potential functional relevance, including convergence between Rousettus and classic echolocators in prestin at a site known to affect hair cell electromotility. We also detected convergence between Yangochiroptera, Rhinolophidea, and oilbirds in TMC1, an important mechanosensory transduction channel in vertebrate hair cells, and observed an amino acid change at the same site within the pore domain. Our results suggest that although most proteins implicated in echolocation in specialized mammals may not have been recruited in birds or Rousettus fruit bats, certain hearing-related loci may have undergone convergent functional changes. Investigating adaptations in diverse echolocators will deepen our understanding of this unusual sensory modality.

A heavyweight early whale pushes the boundaries of vertebrate morphology.

The fossil record of cetaceans documents how terrestrial animals acquired extreme adaptations and transitioned to a fully aquatic lifestyle1,2. In whales, this is associated with a substantial increase in maximum body size. Although an elongate body was acquired early in cetacean evolution3, the maximum body mass of baleen whales reflects a recent diversification that culminated in the blue whale4. More generally, hitherto known gigantism among aquatic tetrapods evolved within pelagic, active swimmers. Here we describe Perucetus colossus-a basilosaurid whale from the middle Eocene epoch of Peru. It displays, to our knowledge, the highest degree of bone mass increase known to date, an adaptation associated with shallow diving5. The estimated skeletal mass of P. colossus exceeds that of any known mammal or aquatic vertebrate. We show that the bone structure specializations of aquatic mammals are reflected in the scaling of skeletal fraction (skeletal mass versus whole-body mass) across the entire disparity of amniotes. We use the skeletal fraction to estimate the body mass of P. colossus, which proves to be a contender for the title of heaviest animal on record. Cetacean peak body mass had already been reached around 30 million years before previously assumed, in a coastal context in which primary productivity was particularly high.

Stuck in a corner: Anthropogenic noise threatens narwhals in their once pristine Arctic habitat.

Niche-conservative species are especially susceptible to changes in their environment, and detecting the negative effects of new stressors in their habitats is vital for safeguarding of these species. In the Arctic, human disturbance including marine traffic and exploration of resources is increasing rapidly due to climate change-induced reduction of sea ice. Here, we show that the narwhal, Monodon monoceros, is extremely sensitive to human-made noise. Narwhals avoided deep diving (> 350 m) with simultaneous reduction of foraging and increased shallow diving activity as a response to either ship sound alone or ship sound with concurrent seismic airgun pulses. Normal behavior decreased by 50 to 75% at distances where received sound levels were below background noise. Narwhals were equally responsive to both disturbance types, hence demonstrating their acute sensitivity to ship sound. This sensitivity coupled with their special behavioral-ecological strategy including a narrow ecological niche and high site fidelity makes them thus especially vulnerable to human impacts in the Arctic.

Toothed whales use distinct vocal registers for echolocation and communication.

Echolocating toothed whales (odontocetes) capture fast-moving prey in dark marine environments, which critically depends on their ability to generate powerful, ultrasonic clicks. How their supposedly air-driven sound source can produce biosonar clicks at depths of >1000 meters, while also producing rich vocal repertoires to mediate complex social communication, remains unknown. We show that odontocetes possess a sound production system based on air driven through nasal passages that is functionally analogous to laryngeal and syringeal sound production. Tissue vibration in different registers produces distinct echolocation and communication signals across all major odontocete clades, and thus provides a physiological basis for classifying their vocal repertoires. The vocal fry register is used by species from porpoises to sperm whales for generating powerful, highly air-efficient echolocation clicks.

Effects of vessel traffic and ocean noise on gray whale stress hormones.

Human use of marinescapes is rapidly increasing, especially in populated nearshore regions where recreational vessel traffic can be dense. Marine animals can have a physiological response to such elevated human activity that can impact individual health and population dynamics. To understand the physiological impacts of vessel traffic on baleen whales, we investigated the adrenal stress response of gray whales (Eschrichtius robustus) to variable vessel traffic levels through an assessment of fecal glucocorticoid metabolite (fGC) concentrations. This analysis was conducted at the individual level, at multiple temporal scales (1-7 days), and accounted for factors that may confound fGC: sex, age, nutritional status, and reproductive state. Data were collected in Oregon, USA, from June to October of 2016-2018. Results indicate significant correlations between fGC, month, and vessel counts from the day prior to fecal sample collection. Furthermore, we show a significant positive correlation between vessel traffic and underwater ambient noise levels, which indicates that noise produced by vessel traffic may be a causal factor for the increased fGC. This study increases knowledge of gray whale physiological response to vessel traffic and may inform management decisions regarding regulations of vessel traffic activities and thresholds near critical whale habitats.

Seismic surveys near gray whale feeding areas off Sakhalin Island, Russia: assessing impact and mitigation effectiveness.

In 2015, two oil and gas companies conducted seismic surveys along the northeast coast of Sakhalin Island, Russia, near western gray whale (Eschrichtius robustus) feeding areas. This population of whales was listed as Critically Endangered at the time of the operations described here but has been reclassified as Endangered since 2018. The number and duration of the 2015 seismic surveys surpassed the level of previous seismic survey activity in this area, elevating concerns regarding disturbance of feeding gray whales and the potential for auditory injury. Exxon Neftegas Limited (ENL) developed a mitigation approach to address these concerns and, more importantly, implemented a comprehensive data collection strategy to assess the effectiveness of this approach. The mitigation approach prioritized completion of the seismic surveys closest to the nearshore feeding area as early in the season as possible, when fewer gray whales would be present. This was accomplished by increasing operational efficiency through the use of multiple seismic vessels and by establishing zones with specific seasonal criteria determining when air gun shutdowns would be implemented. These zones and seasonal criteria were based on pre-season modeled acoustic footprints of the air gun array and on gray whale distribution data collected over the previous 10 years. Real-time acoustic and whale sighting data were instrumental in the implementation of air gun shutdowns. The mitigation effectiveness of these shutdowns was assessed through analyzing short-term behavioral responses and shifts in gray whale distribution due to sound exposure. The overall mitigation strategy of an early survey completion was assessed through bioenergetics models that predict how reduced foraging activity might affect gray whale reproduction and maternal survival. This assessment relied on a total of 17 shore-based and 5 vessel-based teams collecting behavior, distribution, photo-identification, prey, and acoustic data. This paper describes the mitigation approach, the implementation of mitigation measures using real-time acoustic and gray whale location data, and the strategy to assess impacts and mitigation effectiveness.

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.

Acoustic monitoring and analyses of air gun, pile driving, vessel, and ambient sounds during the 2015 seismic surveys on the Sakhalin shelf.

During the summer of 2015, four 4D seismic surveys were conducted on the northeastern Sakhalin shelf near the feeding grounds of the Korean-Okhotsk (western) gray whale (Eschrichtius robustus) population. In addition to the seismic surveys, onshore pile driving activities and vessel operations occurred. Forty autonomous underwater acoustic recorders provided data in the 2 Hz to15 kHz frequency band. Recordings were analyzed to evaluate the characteristics of impulses propagating from the seismic sources. Acoustic metrics analyzed comprised peak sound pressure level (PK), mean square sound pressure level (SPL), sound exposure level (SEL), T100%, T90% (the time intervals that contain the full and 90% of the energy of the impulse), and kurtosis. The impulses analyzed differed significantly due to the variability and complexity of propagation in the shallow water of the northeast Sakhalin shelf. At larger ranges, a seismic precursor propagated in the seabed ahead of the acoustic impulse, and the impulses often interfered with each other, complicating analyses. Additional processing of recordings allowed evaluation and documentation of relevant metrics for pile driving, vessel sounds, and ambient background levels. The computed metrics were used to calibrate acoustic models, generating time resolved estimates of the acoustic levels from seismic surveys, pile driving, and vessel operations on a gray whale distribution grid and along observed gray whale tracks. This paper describes the development of the metrics and the calibrated acoustic models, both of which will be used in work quantifying gray whale behavioral and distribution responses to underwater sounds and to determine whether these observed responses have the potential to impact important parameters at the population level (e.g., reproductive success).

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.

Real-time acoustic monitoring with telemetry to mitigate potential effects of seismic survey sounds on marine mammals: a case study offshore Sakhalin Island.

Exxon Neftegas Ltd. (ENL) carried out three 4D seismic surveys during the summer of 2015. Seismic operations in two of these fields (Odoptu and Chayvo) ensonified the nearshore feeding area of Korean-Okhotsk (western) gray whales (Eschrichtius robustus), potentially disturbing feeding activities. Following model-based optimization of the source design to minimize its lateral acoustic footprint, pre-season modeling was used to compute the acoustic exposure along each survey line. Real-time acoustic data facilitated implementation of mitigation measures aimed to minimize disturbance of whales. Acoustic data originated from underwater recorders deployed on the seafloor. Two complementary approaches were used to transmit recorded sound data to a computer housed at the Central Post (CP), where decisions regarding mitigation shut downs were made. In the first approach, a limited bandwidth (2-2000 Hz) sampling of the data was transmitted via cable to a surface buoy, which relayed these data to a shore station up to 15 km away via digital VHF telemetry. At the shore station, acoustic impulses from the seismic surveys were processed to compute impulse characteristics in the form of estimates of sound exposure level and peak sound pressure level, as well as one-minute-average 1/3-octave power spectral density coefficients, which were then transmitted to the CP via the internet. In the second, the pulse characteristics were computed through algorithms running on an onboard processor in each recorder's surface buoy and sent directly to the CP computer via an Iridium satellite uplink. Both methods of data transfer proved viable, but Iridium transmission achieved the goal without the need for any shore based relay stations and is therefore more operationally efficient than VHF transmission. At the CP, analysts used the real-time acoustic data to calibrate and adjust the output of pre-season acoustical model runs. The acoustic footprint for the active seismic source, advancing synchronously with the motion of the seismic vessel and changing as the sound propagation environment changed, was computed from the calibrated and adjusted model output and integrated through the software Pythagoras with locations of gray whales provided by shore-based observers. This enabled analysts to require air gun array shutdowns before whales were exposed to mean square sound pressure levels greater than the behavioral response threshold of 163 dB re 1 µPa2. The method described here provides a realistic means of mitigating the possible effects of air guns at a behavioral response level, whereas most seismic surveys rely on pre-established mitigation radii to manage the risk of injury to a whale.

Recasting the whale's wonderful net.

Modification of cerebral vasculature helps to cushion the brains of whales and dolphins against injury.

Ancient whale rhodopsin reconstructs dim-light vision over a major evolutionary transition: Implications for ancestral diving behavior.

Cetaceans are fully aquatic mammals that descended from terrestrial ancestors, an iconic evolutionary transition characterized by adaptations for underwater foraging via breath-hold diving. Although the evolutionary history of this specialized behavior is challenging to reconstruct, coevolving sensory systems may offer valuable clues. The dim-light visual pigment, rhodopsin, which initiates phototransduction in the rod photoreceptors of the eye, has provided insight into the visual ecology of depth in several aquatic vertebrate lineages. Here, we use ancestral sequence reconstruction and protein resurrection experiments to quantify light-activation metrics in rhodopsin pigments from ancestors bracketing the cetacean terrestrial-to-aquatic transition. By comparing multiple reconstruction methods on a broadly sampled cetartiodactyl species tree, we generated highly robust ancestral sequence estimates. Our experimental results provide direct support for a blue-shift in spectral sensitivity along the branch separating cetaceans from terrestrial relatives. This blue-shift was 14 nm, resulting in a deep-sea signature (λmax = 486 nm) similar to many mesopelagic-dwelling fish. We also discovered that the decay rates of light-activated rhodopsin increased in ancestral cetaceans, which may indicate an accelerated dark adaptation response typical of deeper-diving mammals. Because slow decay rates are thought to help sequester cytotoxic photoproducts, this surprising result could reflect an ecological trade-off between rod photoprotection and dark adaptation. Taken together, these ancestral shifts in rhodopsin function suggest that some of the first fully aquatic cetaceans could dive into the mesopelagic zone (>200 m). Moreover, our reconstructions indicate that this behavior arose before the divergence of toothed and baleen whales.

Isotopic composition of the eastern gray whale epidermis indicates contribution of prey outside Arctic feeding grounds.

Eastern gray whales' distribution range and plasticity in feeding behavior complicates the understanding of critical life-history such as pregnancy and lactation. Our goal was to determine if females who experienced gestation, gave birth, and lactated their calves, assimilated a high proportion of benthic amphipods from the Bering Sea, which are considered the species' main prey. We used Bayesian stable isotope mixing models to estimate the probability of contribution of food items sampled along the species' distributional range, using isotopic data on amphipods from the Bering Sea, mysids from Vancouver Island, and amphipods and polychaetes from Ojo de Liebre Lagoon. We sampled epidermal tissue from lactating females (n = 25) and calves (n = 34) and analyzed their carbon and nitrogen isotopic composition. Model outcome indicated that benthic amphipods from the Bering Sea were not the primary food for the eastern gray whale. Each mother performed a different feeding strategy, and prey from Vancouver Island were generally as important as that from the Bering Sea. Moreover, model results indicate a constant use of Ojo de Liebre Lagoon as a feeding ground. Our results appear to agree with previous studies that report continuous feeding by females to satisfy certain physiological requirements (e.g., fatty acids omega-6) during migration and breeding time. Future investigations of the isotopic composition of all those prey items that could be assimilated by the eastern gray whale emerge as critical. Both historical and recent information, that would provide insights in the species feeding ecology under past and present environmental conditions, should be considered as equally important to establish conservation and management plans.