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.

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.

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.

Baleen whale prey consumption based on high-resolution foraging measurements.

Baleen whales influence their ecosystems through immense prey consumption and nutrient recycling1-3. It is difficult to accurately gauge the magnitude of their current or historic ecosystem role without measuring feeding rates and prey consumed. To date, prey consumption of the largest species has been estimated using metabolic models3-9 based on extrapolations that lack empirical validation. Here, we used tags deployed on seven baleen whale (Mysticeti) species (n = 321 tag deployments) in conjunction with acoustic measurements of prey density to calculate prey consumption at daily to annual scales from the Atlantic, Pacific, and Southern Oceans. Our results suggest that previous studies3-9 have underestimated baleen whale prey consumption by threefold or more in some ecosystems. In the Southern Ocean alone, we calculate that pre-whaling populations of mysticetes annually consumed 430 million tonnes of Antarctic krill (Euphausia superba), twice the current estimated total biomass of E. superba10, and more than twice the global catch of marine fisheries today11. Larger whale populations may have supported higher productivity in large marine regions through enhanced nutrient recycling: our findings suggest mysticetes recycled 1.2 × 104 tonnes iron yr-1 in the Southern Ocean before whaling compared to 1.2 × 103 tonnes iron yr-1 recycled by whales today. The recovery of baleen whales and their nutrient recycling services2,3,7 could augment productivity and restore ecosystem function lost during 20th century whaling12,13.

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.

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.

Modeling individual growth reveals decreasing gray whale body length and correlations with ocean climate indices at multiple scales.

Changes in body size have been documented across taxa in response to human activities and climate change. Body size influences many aspects of an individual's physiology, behavior, and ecology, ultimately affecting life history performance and resilience to stressors. In this study, we developed an analytical approach to model individual growth patterns using aerial imagery collected via drones, which can be used to investigate shifts in body size in a population and the associated drivers. We applied the method to a large morphological dataset of gray whales (Eschrichtius robustus) using a distinct foraging ground along the NE Pacific coast, and found that the asymptotic length of these whales has declined since around the year 2000 at an average rate of 0.05-0.12 m/y. The decline has been stronger in females, which are estimated to be now comparable in size to males, minimizing sexual dimorphism. We show that the decline in asymptotic length is correlated with two oceanographic metrics acting as proxies of habitat quality at different scales: the mean Pacific Decadal Oscillation index, and the mean ratio between upwelling intensity in a season and the number of relaxation events. These results suggest that the decline in gray whale body size may represent a plastic response to changing environmental conditions. Decreasing body size could have cascading effects on the population's demography, ability to adjust to environmental changes, and ecological influence on the structure of their community. This finding adds to the mounting evidence that body size is shrinking in several marine populations in association with climate change and other anthropogenic stressors. Our modeling approach is broadly applicable across multiple systems where morphological data on megafauna are collected using drones.

Beaked whales and state-dependent decision-making: how does body condition affect the trade-off between foraging and predator avoidance?

Body condition is central to how animals balance foraging with predator avoidance-a trade-off that fundamentally affects animal fitness. Animals in poor condition may accept greater predation risk to satisfy current foraging 'needs', while those in good condition may be more risk averse to protect future 'assets'. These state-dependent behavioural predictions can help interpret responses to human activities, but are little explored in marine animals. This study investigates the influence of body condition on how beaked whales trade-off foraging and predator avoidance. Body density (indicating lipid-energy stores) was estimated for 15 foraging northern bottlenose whales tagged near Jan Mayen, Norway. Composite indices of foraging (diving and echolocation clicks) and anti-predation (long ascents, non-foraging dives and silent periods reducing predator eavesdropping) were negatively related. Experimental sonar exposures led to decreased foraging and increased risk aversion, confirming a foraging/perceived safety trade-off. However, lower lipid stores were not related to a decrease in predator avoidance versus foraging, i.e. worse condition animals did not prioritize foraging. Individual differences (personalities) or reproductive context could offer alternative explanations for the observed state-behaviour relationships. This study provides evidence of foraging/predator-avoidance trade-offs in a marine top predator and demonstrates that animals in worse condition might not always take more risks.

Strong and lasting impacts of past global warming on baleen whales and their prey.

Global warming is affecting the population dynamics and trophic interactions across a wide range of ecosystems and habitats. Translating these real-time effects into their long-term consequences remains a challenge. The rapid and extreme warming period that occurred after the Last Glacial Maximum (LGM) during the Pleistocene-Holocene transition (7-12 thousand years ago) provides an opportunity to gain insights into the long-term responses of natural populations to periods with global warming. The effects of this post-LGM warming period have been assessed in many terrestrial taxa, whereas insights into the impacts of rapid global warming on marine taxa remain limited, especially for megafauna. In order to understand how large-scale climate fluctuations during the post-LGM affected baleen whales and their prey, we conducted an extensive, large-scale analysis of the long-term effects of the post-LGM warming on abundance and inter-ocean connectivity in eight baleen whale and seven prey (fish and invertebrates) species across the Southern and the North Atlantic Ocean; two ocean basins that differ in key oceanographic features. The analysis was based upon 7032 mitochondrial DNA sequences as well as genome-wide DNA sequence variation in 100 individuals. The estimated temporal changes in genetic diversity during the last 30,000 years indicated that most baleen whale populations underwent post-LGM expansions in both ocean basins. The increase in baleen whale abundance during the Holocene was associated with simultaneous changes in their prey and climate. Highly correlated, synchronized and exponential increases in abundance in both baleen whales and their prey in the Southern Ocean were indicative of a dramatic increase in ocean productivity. In contrast, the demographic fluctuations observed in baleen whales and their prey in the North Atlantic Ocean were subtle, varying across taxa and time. Perhaps most important was the observation that the ocean-wide expansions and decreases in abundance that were initiated by the post-LGM global warming, continued for millennia after global temperatures stabilized, reflecting persistent, long-lasting impacts of global warming on marine fauna.

Sowerby's beaked whale biosonar and movement strategy indicate deep-sea foraging niche differentiation in mesoplodont whales.

Closely related species are expected to diverge in foraging strategy, reflecting the evolutionary drive to optimize foraging performance. The most speciose cetacean genus, Mesoplodon, comprises beaked whales with little diversity in external morphology or diet, and overlapping distributions. Moreover, the few studied species of beaked whales (Ziphiidae) show very similar foraging styles with slow, energy-conserving movement during long, deep foraging dives. This raises the question of what factors drive their speciation. Using data from animal-attached tags and aerial imagery, we tested the hypothesis that two similar-sized mesoplodonts, Sowerby's (Mesoplodon bidens) and Blainville's (Mesoplodon densirostris) beaked whales, exploit a similar low-energy niche. We show that, compared with the low-energy strategist Blainville's beaked whale, Sowerby's beaked whale lives in the fast lane. While targeting a similar mesopelagic/bathypelagic foraging zone, they consistently swim and hunt faster, perform shorter deep dives, and echolocate at a faster rate with higher frequency clicks. Further, extensive near-surface travel between deep dives challenges the interpretation of beaked whale shallow inter-foraging dives as a management strategy for decompression sickness. The distinctively higher frequency echolocation clicks do not hold apparent foraging benefits. Instead, we argue that a high-speed foraging style influences dive duration and echolocation behaviour, enabling access to a distinct prey population. Our results demonstrate that beaked whales exploit a broader diversity of deep-sea foraging and energetic niches than hitherto suspected. The marked deviation of Sowerby's beaked whales from the typical ziphiid foraging strategy has potential implications for their response to anthropogenic sounds, which appears to be strongly behaviourally driven in other ziphiids.

A machine learning pipeline for classification of cetacean echolocation clicks in large underwater acoustic datasets.

Machine learning algorithms, including recent advances in deep learning, are promising for tools for detection and classification of broadband high frequency signals in passive acoustic recordings. However, these methods are generally data-hungry and progress has been limited by challenges related to the lack of labeled datasets adequate for training and testing. Large quantities of known and as yet unidentified broadband signal types mingle in marine recordings, with variability introduced by acoustic propagation, source depths and orientations, and interacting signals. Manual classification of these datasets is unmanageable without an in-depth knowledge of the acoustic context of each recording location. A signal classification pipeline is presented which combines unsupervised and supervised learning phases with opportunities for expert oversight to label signals of interest. The method is illustrated with a case study using unsupervised clustering to identify five toothed whale echolocation click types and two anthropogenic signal categories. These categories are used to train a deep network to classify detected signals in either averaged time bins or as individual detections, in two independent datasets. Bin-level classification achieved higher overall precision (>99%) than click-level classification. However, click-level classification had the advantage of providing a label for every signal, and achieved higher overall recall, with overall precision from 92 to 94%. The results suggest that unsupervised learning is a viable solution for efficiently generating the large, representative training sets needed for applications of deep learning in passive acoustics.

Discrete-space continuous-time models of marine mammal exposure to Navy sonar.

Assessing the patterns of wildlife attendance to specific areas is relevant across many fundamental and applied ecological studies, particularly when animals are at risk of being exposed to stressors within or outside the boundaries of those areas. Marine mammals are increasingly being exposed to human activities that may cause behavioral and physiological changes, including military exercises using active sonars. Assessment of the population-level consequences of anthropogenic disturbance requires robust and efficient tools to quantify the levels of aggregate exposure for individuals in a population over biologically relevant time frames. We propose a discrete-space, continuous-time approach to estimate individual transition rates across the boundaries of an area of interest, informed by telemetry data collected with uncertainty. The approach allows inferring the effect of stressors on transition rates, the progressive return to baseline movement patterns, and any difference among individuals. We apply the modeling framework to telemetry data from Blainville's beaked whale (Mesoplodon densirostris) tagged in the Bahamas at the Atlantic Undersea Test and Evaluation Center (AUTEC), an area used by the U.S. Navy for fleet readiness training. We show that transition rates changed as a result of exposure to sonar exercises in the area, reflecting an avoidance response. Our approach supports the assessment of the aggregate exposure of individuals to sonar and the resulting population-level consequences. The approach has potential applications across many applied and fundamental problems where telemetry data are used to characterize animal occurrence within specific areas.

Vast assembly of vocal marine mammals from diverse species on fish spawning ground.

Observing marine mammal (MM) populations continuously in time and space over the immense ocean areas they inhabit is challenging but essential for gathering an unambiguous record of their distribution, as well as understanding their behaviour and interaction with prey species. Here we use passive ocean acoustic waveguide remote sensing (POAWRS) in an important North Atlantic feeding ground to instantaneously detect, localize and classify MM vocalizations from diverse species over an approximately 100,000 km(2) region. More than eight species of vocal MMs are found to spatially converge on fish spawning areas containing massive densely populated herring shoals at night-time and diffuse herring distributions during daytime. We find the vocal MMs divide the enormous fish prey field into species-specific foraging areas with varying degrees of spatial overlap, maintained for at least two weeks of the herring spawning period. The recorded vocalization rates are diel (24 h)-dependent for all MM species, with some significantly more vocal at night and others more vocal during the day. The four key baleen whale species of the region: fin, humpback, blue and minke have vocalization rate trends that are highly correlated to trends in fish shoaling density and to each other over the diel cycle. These results reveal the temporospatial dynamics of combined multi-species MM foraging activities in the vicinity of an extensive fish prey field that forms a massive ecological hotspot, and would be unattainable with conventional methodologies. Understanding MM behaviour and distributions is essential for management of marine ecosystems and for accessing anthropogenic impacts on these protected marine species.

Isotopes from fossil coronulid barnacle shells record evidence of migration in multiple Pleistocene whale populations.

Migration is an integral feature of modern mysticete whale ecology, and the demands of migration may have played a key role in shaping mysticete evolutionary history. Constraining when migration became established and assessing how it has changed through time may yield valuable insight into the evolution of mysticete whales and the oceans in which they lived. However, there are currently few data which directly assess prehistoric mysticete migrations. Here we show that calcite δ18O profiles of two species of modern whale barnacles (coronulids) accurately reflect the known migration routes of their host whales. We then analyze well-preserved fossil coronulids from three different locations along the eastern Pacific coast, finding that δ18O profiles from these fossils exhibit trends and ranges similar to modern specimens. Our results demonstrate that migration is an ancient behavior within the humpback and gray whale lineages and that multiple Pleistocene populations were undertaking migrations of an extent similar to those of the present day.

A functional enrichment test for molecular convergent evolution finds a clear protein-coding signal in echolocating bats and whales.

Distantly related species entering similar biological niches often adapt by evolving similar morphological and physiological characters. How much genomic molecular convergence (particularly of highly constrained coding sequence) contributes to convergent phenotypic evolution, such as echolocation in bats and whales, is a long-standing fundamental question. Like others, we find that convergent amino acid substitutions are not more abundant in echolocating mammals compared to their outgroups. However, we also ask a more informative question about the genomic distribution of convergent substitutions by devising a test to determine which, if any, of more than 4,000 tissue-affecting gene sets is most statistically enriched with convergent substitutions. We find that the gene set most overrepresented (q-value = 2.2e-3) with convergent substitutions in echolocators, affecting 18 genes, regulates development of the cochlear ganglion, a structure with empirically supported relevance to echolocation. Conversely, when comparing to nonecholocating outgroups, no significant gene set enrichment exists. For aquatic and high-altitude mammals, our analysis highlights 15 and 16 genes from the gene sets most affected by molecular convergence which regulate skin and lung physiology, respectively. Importantly, our test requires that the most convergence-enriched set cannot also be enriched for divergent substitutions, such as in the pattern produced by inactivated vision genes in subterranean mammals. Showing a clear role for adaptive protein-coding molecular convergence, we discover nearly 2,600 convergent positions, highlight 77 of them in 3 organs, and provide code to investigate other clades across the tree of life.

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).

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.

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.

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.