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Dichromatic color vision is mediated by two cone visual pigments in many eutherian mammals. After reentry into the sea, early cetaceans lost their violet-sensitive visual pigment (short wavelength-sensitive 1) independently in the baleen and toothed whale ancestors and thus obtained only monochromatic cone vision. Subsequently, losses of the middle/long wavelength-sensitive (M/LWS) pigment have also been reported in multiple whale lineages, leading to rhodopsin (RH1)-mediated rod monochromatic vision. To further elucidate the phenotypic evolution of whale visual pigments, we assessed the spectral tuning of both M/LWS and RH1 from representative cetacean taxa. Interestingly, although the coding sequences for M/LWS are intact in both the pygmy right whale and the Baird's beaked whale, no spectral sensitivity was detected in vitro. Pseudogenization of other cone vision-related genes is observed in the pygmy right whale, suggesting a loss of cone-mediated vision. After ancestral sequence reconstructions, ancient M/LWS pigments from cetacean ancestors were resurrected and functionally measured. Spectral tuning of M/LWS from the baleen whale ancestor shows that it is green sensitive, with a 40-nm shift in sensitivity to a shorter wavelength. For the ancestor of sperm whales, although no spectral sensitivity could be recorded for its M/LWS pigment, a substantial sensitivity shift (20 to 30 nm) to a shorter wavelength may have also occurred before its functional inactivation. The parallel phenotypic evolution of M/LWS to shorter wavelength sensitivity might be visual adaptations in whales allowing more frequent deep-sea activities, although additional ecological differentiations may have led to their subsequent losses., Competing Interests: Conflict of Interest There is no conflict of interest., (© The Author(s) 2024. Published by Oxford University Press on behalf of Society for Molecular Biology and Evolution.)

Background: Habitat transitions have considerable consequences in organism homeostasis, as they require the adjustment of several concurrent physiological compartments to maintain stability and adapt to a changing environment. Within the range of molecules with a crucial role in the regulation of different physiological processes, neuropeptides are key agents. Here, we examined the coding status of several neuropeptides and their receptors with pleiotropic activity in Cetacea., Results: Analysis of 202 mammalian genomes, including 41 species of Cetacea, exposed an intricate mutational landscape compatible with gene sequence modification and loss. Specifically for Cetacea, in the 12 genes analysed we have determined patterns of loss ranging from species-specific disruptive mutations (e.g. neuropeptide FF-amide peptide precursor; NPFF) to complete erosion of the gene across the cetacean stem lineage (e.g. somatostatin receptor 4; SSTR4)., Conclusions: Impairment of some of these neuromodulators may have contributed to the unique energetic metabolism, circadian rhythmicity and diving response displayed by this group of iconic mammals., (© 2024. The Author(s).)

The marine environment presents challenges for wound healing in cetaceans, despite their remarkable recovery abilities with minimal infections or complications. However, the molecular mechanism underlying this efficient wound healing remains underexplored. To better understand the molecular mechanisms behind wound healing in cetaceans, we investigated the evolutionary patterns of 37 wound healing-related genes in representative mammals. We found wound healing-related genes experience adaptive evolution in cetaceans: (1) Three extrinsic coagulation pathway-related genes-tissue factor (F3), coagulation factor VII (F7), and coagulation factor X (F10)-are subject to positive selection in cetaceans, which might promote efficient hemostasis after injury; positive selection in transforming growth factor-beta 2 (TGF-β2), transforming growth factor-beta 3 (TGF-β3), and platelet-derived growth factor D (PDGFD), which play immunological roles in wound healing, may help cetaceans enhance inflammatory response and tissue debridement. (2) Coagulation factor XII (F12) is the initiation factor in the intrinsic coagulation pathway. It had a premature stop codon mutation and was subjected to selective stress relaxation in cetaceans, suggesting that the early termination of F12 may help cetaceans avoid the risk of vascular blockage during diving. (3) Fibrinogen alpha chain (FGA) and FIII, which were detected to contain the specific amino acid substitutions in marine mammals, indicating similar evolutionary mechanisms might exist among marine mammals to maintain strong wound-healing ability. Thus, our research provides further impetus to study the evolution of the wound healing system in cetaceans and other marine mammals, extending knowledge of preventing coagulation disorder and atherosclerosis in humans., (© 2023 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

Cetacean reinvasion of the aquatic realm is an iconic ecological transition that led to drastic modifications of the mammalian body plan, especially in the axial skeleton. Relative to the vertebral column of other mammals that is subdivided into numerous anatomical regions, regional boundaries of the cetacean backbone appear obscured. Whether the traditional mammalian regions are present in cetaceans but hard to detect due to anatomical homogenization or if regions have been entirely repatterned remains unresolved. Here we combine a segmented linear regression approach with spectral clustering to quantitatively investigate the number, position, and homology of vertebral regions across 62 species from all major cetacean clades. We propose the Nested Regions hypothesis under which the cetacean backbone is composed of six homologous modules subdivided into six to nine post-cervical regions, with the degree of regionalization dependent on vertebral count and ecology. Compared to terrestrial mammals, the cetacean backbone is less regionalized in the precaudal segment but more regionalized in the caudal segment, indicating repatterning of the vertebral column associated with the transition from limb-powered to axial-driven locomotion., (© 2024. The Author(s).)

Cetaceans represent a natural experiment within the tree of life in which a lineage changed from terrestrial to aquatic habitats. This shift involved phenotypic modifications, representing an opportunity to explore the genetic bases of phenotypic diversity. Among the different molecular systems that maintain cellular homeostasis, ion channels are crucial for the proper physiological functioning of all living species. This study aims to explore the evolution of ion channels during the evolutionary history of cetaceans. To do so, we created a bioinformatic pipeline to annotate the repertoire of ion channels in the genome of the species included in our sampling. Our main results show that cetaceans have, on average, fewer protein-coding genes and a higher percentage of annotated ion channels than non-cetacean mammals. Signals of positive selection were detected in ion channels related to the heart, locomotion, visual and neurological phenotypes. Interestingly, we predict that the NaV1.5 ion channel of most toothed whales (odontocetes) is sensitive to tetrodotoxin, similar to NaV1.7, given the presence of tyrosine instead of cysteine, in a specific position of the ion channel. Finally, the gene turnover rate of the cetacean crown group is more than three times faster than that of non-cetacean mammals., (© 2024. The Author(s).)

While rhythm can facilitate and enhance many aspects of behavior, its evolutionary trajectory in vocal communication systems remains enigmatic. We can trace evolutionary processes by investigating rhythmic abilities in different species, but research to date has largely focused on songbirds and primates. We present evidence that cetaceans-whales, dolphins, and porpoises-are a missing piece of the puzzle for understanding why rhythm evolved in vocal communication systems. Cetaceans not only produce rhythmic vocalizations but also exhibit behaviors known or thought to play a role in the evolution of different features of rhythm. These behaviors include vocal learning abilities, advanced breathing control, sexually selected vocal displays, prolonged mother-infant bonds, and behavioral synchronization. The untapped comparative potential of cetaceans is further enhanced by high interspecific diversity, which generates natural ranges of vocal and social complexity for investigating various evolutionary hypotheses. We show that rhythm (particularly isochronous rhythm, when sounds are equally spaced in time) is prevalent in cetacean vocalizations but is used in different contexts by baleen and toothed whales. We also highlight key questions and research areas that will enhance understanding of vocal rhythms across taxa. By coupling an infraorder-level taxonomic assessment of vocal rhythm production with comparisons to other species, we illustrate how broadly comparative research can contribute to a more nuanced understanding of the prevalence, evolution, and possible functions of rhythm in animal communication., Competing Interests: Competing interests statement:The authors declare no competing interest.

An underwater acoustic recorder was moored off Heard Island from September 2017 through March 2018 to listen for marine mammals. Analysis of data was initially conducted by visual inspection of long-term spectral averages to reveal sounds from sperm whales, Antarctic and pygmy blue whales, fin whales, minke whales, odontocete whistles, and noise from nearby ships. Automated detection of sperm whale clicks revealed they were seldom detected from September through January (n = 35 h) but were detected nearly every day of February and March (n = 684 h). Additional analysis of these detections revealed further diel and demographic patterns., (© 2024 Author(s). All article content, except where otherwise noted, is licensed under a Creative Commons Attribution (CC BY) license (https://creativecommons.org/licenses/by/4.0/).)

The secondary adaptation of Cetacea to a fully marine lifestyle raises the question of their ability to maintain their water balance in a hyperosmotic environment. Cetacea have access to four potential sources of water: surrounding salt oceanic water, dietary free water, metabolic water and inhaled water vapour to a lesser degree. Here, we measured the 18O/16O oxygen isotope ratio of blood plasma from 13 specimens belonging to two species of Cetacea raised under human care (four killer whales Orcinus orca, nine common bottlenose dolphins Tursiops truncatus) to investigate and quantify the contribution of preformed water (dietary free water, surrounding salt oceanic water) and metabolic water to Cetacea body water using a box-modelling approach. The oxygen isotope composition of Cetacea blood plasma indicates that dietary free water and metabolic water contribute to more than 90% of the total water input in weight for cetaceans, with the remaining 10% consisting of inhaled water vapour and surrounding water accidentally ingested or absorbed through the skin. Moreover, the contribution of metabolic water appears to be more important in organisms with a more lipid-rich diet. Beyond these physiological and conservation biology implications, this study opens up questions that need to be addressed, such as the applicability of the oxygen isotope composition of cetacean body fluids and skeletal elements as an environmental proxy of the oxygen isotope composition of present and past marine waters., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Global reductions in the underwater radiated noise levels from cargo vessels are needed to reduce increasing cumulative impacts to marine wildlife. We use a vessel exposure simulation model to examine how reducing vessel source levels through slowdowns and technological modifications can lessen impacts on marine mammals. We show that the area exposed to ship noise reduces markedly with moderate source-level reductions that can be readily achieved with small reductions in speed. Moreover, slowdowns reduce all impacts to marine mammals despite the longer time that a slower vessel takes to pass an animal. We conclude that cumulative noise impacts from the global fleet can be reduced immediately by slowdowns. This solution requires no modification to ships and is scalable from local speed reductions in sensitive areas to ocean basins. Speed reductions can be supplemented by routing vessels away from critical habitats and by technological modifications to reduce vessel noise.

Toothed whales evolved a third way of making sounds similar to that of land mammals and birds.

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.

Long-term monitoring datasets are fundamental to understand physical and ecological responses to environmental changes, supporting management and conservation. The data should be reliable, with the sources of bias identified and quantified. CETUS Project is a cetacean monitoring programme in the Eastern North Atlantic, based on visual methods of data collection. This study aims to assess data quality and bias in the CETUS dataset, by 1) applying validation methods, through photographic confirmation of species identification; 2) creating data quality criteria to evaluate the observer's experience; and 3) assessing bias to the number of sightings collected and to the success in species identification. Through photographic validation, the species identification of 10 sightings was corrected and a new species was added to the CETUS dataset. The number of sightings collected was biased by external factors, mostly by sampling effort but also by weather conditions. Ultimately, results highlight the importance of identifying and quantifying data bias, while also yielding guidelines for data collection and processing, relevant for species monitoring programmes based on visual methods., (© 2022. The Author(s).)

Cetaceans have massive vascular plexuses (retia mirabilia) whose function is unknown. All cerebral blood flow passes through these retia, and we hypothesize that they protect cetacean brains from locomotion-generated pulsatile blood pressures. We propose that cetaceans have evolved a pulse-transfer mechanism that minimizes pulsatility in cerebral arterial-to-venous pressure differentials without dampening the pressure pulses themselves. We tested this hypothesis using a computational model based on morphology from 11 species and found that the large arterial capacitance in the retia, coupled with the small extravascular capacitance in the cranium and vertebral canal, could protect the cerebral vasculature from 97% of systemic pulsatility. Evolution of the retial complex in cetaceans-likely linked to the development of dorsoventral fluking-offers a distinctive solution to adverse locomotion-generated vascular pulsatility.

Body sizes of marine amniotes span six orders of magnitude, yet the factors that governed the evolution of this diversity are largely unknown. High primary production of modern oceans is considered a prerequisite for the emergence of cetacean giants, but that condition cannot explain gigantism in Triassic ichthyosaurs. We describe the new giant ichthyosaur Cymbospondylus youngorum sp. nov. with a 2-meter-long skull from the Middle Triassic Fossil Hill Fauna of Nevada, USA, underscoring rapid size evolution despite the absence of many modern primary producers. Surprisingly, the Fossil Hill Fauna rivaled the composition of modern marine mammal faunas in terms of size range, and energy-flux models suggest that Middle Triassic marine food webs were able to support several large-bodied ichthyosaurs at high trophic levels, shortly after ichthyosaur origins.

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., Competing Interests: The author has declared that no competing interests exist.

Bottlenose dolphins are a behaviorally complex, social species that display a variety of social behaviors. Because of this, it is important for zoological facilities to strive to ensure animals display species-appropriate levels of social behavior. The current study is part of the multi-institutional study entitled "Towards understanding the welfare of cetaceans in zoos and aquariums" commonly referred to as the Cetacean Welfare Study. All participating facilities were accredited by the Alliance of Marine Mammal Parks and Aquariums and/or the Association of Zoos and Aquariums. Behavioral data were collected on 47 bottlenose dolphins representing two subspecies, Tursiops truncatus and Tursiops aduncus, at 25 facilities. The social behaviors of group related activity (group active) as well as interacting with conspecifics (interact with conspecific) were examined for their relationships to both animal management factors and habitat characteristics. The behavioral state of group active and the rate of interact with conspecific were both positively related to the frequency of receiving new forms of environmental enrichment. Both were inversely related to the random scheduling of environmental enrichment. Additional results suggested interact with conspecific was inversely related with daytime spatial experience and that males displayed group active more than females. Overall, the results suggested that animal management techniques such as the type and timing of enrichment may be more important to enhance social behavior than habitat characteristics or the size of the habitat. Information gained from this study can help facilities with bottlenose dolphins manage their enrichment programs in relation to social behaviors., Competing Interests: The authors have read the journal’s policy, and the authors of the study have the following competing interests to declare: LM and LL are affiliated with the Chicago Zoological Society – Brookfield Zoo (an AZA and AMMPA accredited zoo), and Chicago Zoological Society provides a salary for LM. Before the study, JM was previously affiliated with Disney’s Animal Kingdom (an AZA accredited zoo). The Seas® Epcot® Walt Disney World® Resort, Dolphin Island – Resorts World Sentosa, and Loro Parque and Loro Parque Fundación provided funding to the Chicago Zoological Society for this study. This does not alter our adherence to PLOS ONE policies on sharing data and materials. There are no patents, products in development or marketed products associated with this research to declare.

Cetaceans are long-lived, social species that are valued as ambassadors inspiring the public to engage in conservation action. Under professional care, they are critical partners with the scientific community to understanding the biology, behavior, physiology, health, and welfare requirements of this taxonomic group. The Cetacean Welfare Study was a highly collaborative research effort among zoos and aquariums accredited by the Alliance for Marine Mammal Parks and Aquariums and/or the Association of Zoos & Aquariums that provided important empirical and comparative information on the care and management of cetaceans. The goal was to identify factors that were related to the welfare of bottlenose dolphins and to develop reference intervals and values for common and novel indicators of health and welfare for common bottlenose dolphins (Tursiops truncatus), Indo-Pacific bottlenose dolphins (Tursiops aduncus), beluga whales (Delphinapterus leucas), and Pacific white-sided dolphins (Lagenorhynchus obliquidens). Data were collected from cetaceans at 43 accredited zoos and aquariums in seven countries in 2018 and 2019. This overview presents a summary of findings from the initial research articles that resulted from the study titled "Towards understanding the welfare of cetaceans in zoos and aquariums." With multiple related objectives, animal-based metrics were used to advance frameworks of clinical care and target key conditions that were associated with good welfare of cetaceans in zoo and aquarium environments. As a result of this collaboration, species-specific reference intervals and values for blood variables and fecal hormone metabolites were developed and are freely available in an iOS application called ZooPhysioTrak. The results suggested that environmental enrichment programs and social management factors were more strongly related to behaviors likely indicative of positive welfare than habitat characteristics for common and Indo-Pacific bottlenose dolphins. These findings can be widely applied to optimize care and future science-based welfare practice., Competing Interests: The authors have read the journal’s policy, and the authors of the study have the following competing interests to declare: LJM and LKL are affiliated with the Chicago Zoological Society – Brookfield Zoo (an AZA and AMMPA accredited zoo), and Chicago Zoological Society provides a salary for LJM. Before the study, JDM was previously affiliated with Disney’s Animal Kingdom (an AZA accredited zoo). The Seas® Epcot® Walt Disney World® Resort, Dolphin Island – Resorts World Sentosa, and Loro Parque and Loro Parque Fundación provided funding to the Chicago Zoological Society for this study. This does not alter our adherence to PLOS ONE policies on sharing data and materials. There are no patents, products in development or marketed products associated with this research to declare.

Tail flukes as well as the dorsal fin are the apomorphic traits of cetaceans which appeared during the evolutionary process of adaptation to the aquatic life. Both appendages present a wing-like shape associated with lift generation and low drag. We hypothesized that the evolution of fins as lifting structures led to a generic wing design, where the dimensionless parameters of the fin cross-sections are invariant with respect to the body length and taxonomy of small cetaceans (Hypothesis I). We also hypothesized that constraints on variability of a generic fin shape are associated with the primary function of the fin as a fixed or flapping hydrofoil (Hypothesis II). To verify these hypotheses, we examined how the variation in the fin's morphological traits is linked to the primary function, species and body length. Hydrodynamic characteristics of the fin cross-sections were examined with the CFD software and compared with similar engineered airfoils. Generic wing design of both fins was found in a wing-like planform and a streamlined cross-sectional geometry optimized for lift generation. Divergence in a generic fin shape both on the planform and cross-sectional level was found to be related with the fin specialization in fixed or flapping hydrofoil function. Cross-sections of the dorsal fin were found to be optimized for the narrow range of small angles of attack. Cross-sections of tail flukes were found to be more stable for higher angles of attack and had gradual stall characteristics. The obtained results provide an insight into the divergent evolutionary pathways of a generic wing-like shape of the fins of cetaceans under specific demands of thrust production, swimming stability and turning control., Competing Interests: The authors have declared that no competing interests exist.

Plasticity in the cardiac function of a marine mammal facilitates rapid adjustments to the contrasting metabolic demands of breathing at the surface and diving during an extended apnea. By matching their heart rate ( f H ) to their immediate physiological needs, a marine mammal can improve its metabolic efficiency and maximize the proportion of time spent underwater. Respiratory sinus arrhythmia (RSA) is a known modulation of f H that is driven by respiration and has been suggested to increase cardiorespiratory efficiency. To investigate the presence of RSA in cetaceans and the relationship between f H , breathing rate ( f R ) and body mass ( M b ), we measured simultaneous f H and f R in five cetacean species in human care. We found that a higher f R was associated with a higher mean instantaneous f H (i f H ) and minimum i f H of the RSA. By contrast, f H scaled inversely with M b such that larger animals had lower mean and minimum i f H s of the RSA. There was a significant allometric relationship between maximum i f H of the RSA and M b , but not f R , which may indicate that this parameter is set by physical laws and not adjusted dynamically with physiological needs. RSA was significantly affected by f R and was greatly reduced with small increases in f R . Ultimately, these data show that surface f H s of cetaceans are complex and the f H patterns we observed are controlled by several factors. We suggest the importance of considering RSA when interpreting f H measurements and particularly how f R may drive f H changes that are important for efficient gas exchange. This article is part of the theme issue 'Measuring physiology in free-living animals (Part I)'.

Understanding the trophic niches of marine apex predators is necessary to understand interactions between species and to achieve sustainable, ecosystem-based fisheries management. Here, we review the stable carbon and nitrogen isotope ratios for biting marine mammals inhabiting the Atlantic Ocean to test the hypothesis that the relative position of each species within the isospace is rather invariant and that common and predictable patterns of resource partitioning exists because of constrains imposed by body size and skull morphology. Furthermore, we analyze in detail two species-rich communities to test the hypotheses that marine mammals are gape limited and that trophic position increases with gape size. The isotopic niches of species were highly consistent across regions and the topology of the community within the isospace was well conserved across the Atlantic Ocean. Furthermore, pinnipeds exhibited a much lower diversity of isotopic niches than odontocetes. Results also revealed body size as a poor predictor of the isotopic niche, a modest role of skull morphology in determining it, no evidence of gape limitation and little overlap in the isotopic niche of sympatric species. The overall evidence suggests limited trophic flexibility for most species and low ecological redundancy, which should be considered for ecosystem-based fisheries management., (© 2021. The Author(s).)

Scientific information is vital to the conservation of cetaceans and the management of whale-watching activities. The southern coastal waters of Sri Lanka are near a narrow continental shelf and biologically abundant in cetacean species. Although the occurrence of cetaceans has been investigated in certain waters of Sri Lanka, few surveys have been conducted along the southern coast. To fill this gap, we conducted boat-based surveys from January to May 2017 to investigate the occurrence, diversity, and behavior of cetaceans in the waters off Mirissa, covering a survey area of 788.9 km 2 . During 55 survey days, we recorded a total of 242 cetacean sightings and identified at least 9 species (3 mysticetes and 6 odontocetes). The blue whale was the most common mysticete species (167 of 174 mysticete encounters), followed by the Omura's whale (4 of 174) and Bryde's whale (3 of 174). The spinner dolphin was the most common odontocete species (28 of 68 odontocete encounters), followed by the sperm whale (18 of 68), common bottlenose dolphin (13 of 68), short-finned pilot whale (5 of 68), melon-headed whale (2 of 68), and killer whale. Blue whales and sperm whales exhibited a clear preference for outer shelf and high slope areas, and blue whales were observed feeding along these waters. The present study provides near-baseline information on cetacean occurrence and diversity in whale-watching waters off southern Sri Lanka, and highlights the urgent need for proper management strategies for whale-watching activities., (© 2021 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

Animal culture and social bonds are relevant to wildlife conservation because they influence patterns of geography, behavior, and strategies of survival. Numerous examples of socially-driven habitat partitioning and ecological-niche specialization can be found among vertebrates, including toothed whales. But such social-ecological dynamics, described here as 'social niche partitioning', are not known among baleen whales, whose societies-particularly on foraging grounds-are largely perceived as unstructured and incidental to matters of habitat use and conservation. However, through 16 years of behavioral observations and photo-identifications of humpback whales (Megaptera novaeangliae) feeding within a fjord system in the Canadian Pacific (primarily within Gitga'at First Nation waters), we have documented long-term pair bonds (up to 12 years) as well as a complex societal structure, which corresponds closely to persistent patterns in feeding strategy, long-term site fidelity (extended occupancy and annual rate of return up to 75%), specific geographic preferences within the fjord system, and other forms of habitat use. Randomization tests of network congruency and clustering algorithms were used to test for overlap in patterns of social structure and habitat use, which confirmed the occurrence of social niche partitioning on the feeding grounds of this baleen whale species. In addition, we document the extensive practice of group bubble net feeding in Pacific Canada. This coordinated feeding behavior was found to strongly mediate the social structure and habitat use within this humpback whale society. Additionally, during our 2004-2019 study, we observed a shift in social network structure in 2010-2012, which corresponded with environmental and demographic shifts including a sudden decline in the population's calving rate. Our findings indicate that the social lives of humpback whales, and perhaps baleen whales generally, are more complex than previously supposed and should be a primary consideration in the assessment of potential impacts to important habitat., Competing Interests: The authors have declared that no competing interests exist.

The ability to monitor population dynamics and detect major changes in population trend is essential for wildlife conservation and management. However, this is often challenging for cetaceans as surveys typically cover only a portion of a population's range and conventional stock assessment methods cannot then distinguish whether apparent changes in abundance reflect real changes in population size or shifts in distribution. We developed and tested methods for estimating population size and trend and detecting changes in population trend in the context of shifting habitat by integrating additional data into distance-sampling analysis. Previous research has shown that incorporating habitat information can improve population size estimates for highly mobile species with dynamic spatial distributions. Here, using simulated datasets representative of a large whale population, we demonstrate that incorporating individual mark-recapture data can increase the accuracy and precision of trend estimation and the power to distinguish whether apparent changes in abundance reflect changes in population trend or distribution shifts. We recommend that similar simulation studies are conducted for specific cetacean populations to assess the potential for detecting changes in population dynamics given available data. This approach is especially important wherever population change may be confounded with long-term change in distribution patterns associated with regime shifts or climate change., Competing Interests: The authors have declared that no competing interests exist.

The natural endocast Museo di Geologia e Paleontologia of the Università degli Studi di Torino (MGPT)-PU 13873 is described and analyzed in order to interpret its taxonomic affinities and its potential significance on our understanding of cetacean brain evolution. The endocast is from the early Miocene of Piedmont (between ca. 19 and 16 million years ago), Northwestern Italy, and shows a number of plesiomorphic characters. These include: scarcely rounded cerebral hemispheres, cerebellum exposed in dorsal view with little superimposition by the cerebral hemispheres, short temporal lobe, and long sylvian fissure. The distance between the hypophysis and the rostral pons is particularly high, as it was determined by the calculus of the hypothalamus quotient, suggesting that the development of a deep interpeduncular fossa was not as advanced as in living odontocetes. The encephalization quotient (EQ) of MGPT-PU 13873 is ~1.81; therefore, this specimen shows an EQ in line with other fossil whales of the same geological age (early Miocene). Comparative analysis shows that there is a critical lack of data from the late Miocene and Pliocene that prevents us to fully understand the recent evolution of the EQ diversity in whales. Moreover, the past diversity of brain size and shape in mysticetes is virtually unknown. All these observations point to the need of additional efforts to uncover evolutionary patterns and processes on cetacean brain evolution., (© 2020 Wiley Periodicals LLC.)

Remote acquisition of information on ecosystem dynamics is essential for conservation management, especially for the deep ocean. Soundscape offers unique opportunities to study the behavior of soniferous marine animals and their interactions with various noise-generating activities at a fine temporal resolution. However, the retrieval of soundscape information remains challenging owing to limitations in audio analysis techniques that are effective in the face of highly variable interfering sources. This study investigated the application of a seafloor acoustic observatory as a long-term platform for observing marine ecosystem dynamics through audio source separation. A source separation model based on the assumption of source-specific periodicity was used to factorize time-frequency representations of long-duration underwater recordings. With minimal supervision, the model learned to discriminate source-specific spectral features and prove to be effective in the separation of sounds made by cetaceans, soniferous fish, and abiotic sources from the deep-water soundscapes off northeastern Taiwan. Results revealed phenological differences among the sound sources and identified diurnal and seasonal interactions between cetaceans and soniferous fish. The application of clustering to source separation results generated a database featuring the diversity of soundscapes and revealed a compositional shift in clusters of cetacean vocalizations and fish choruses during diurnal and seasonal cycles. The source separation model enables the transformation of single-channel audio into multiple channels encoding the dynamics of biophony, geophony, and anthropophony, which are essential for characterizing the community of soniferous animals, quality of acoustic habitat, and their interactions. Our results demonstrated the application of source separation could facilitate acoustic diversity assessment, which is a crucial task in soundscape-based ecosystem monitoring. Future implementation of soundscape information retrieval in long-term marine observation networks will lead to the use of soundscapes as a new tool for conservation management in an increasingly noisy ocean., Competing Interests: The authors have declared that no competing interests exist.

The ancestors of Cetacea underwent profound morpho-physiological alterations. By displaying an exclusive aquatic existence, cetaceans evolved unique patterns of locomotor activity, vigilant behaviour, thermoregulation and circadian rhythmicity. Deciphering the molecular landscape governing many of these adaptations is key to understand the evolution of phenotypes. Here, we investigate Cortistatin (CORT), a neuropeptide displaying an important role in mammalian biorhythm regulation. This neuropeptide is a known neuroendocrine factor, stimulating slow-wave sleep, but also involved in the regulation of energy metabolism and hypomotility inducement. We assessed the functional status of CORT in 359 mammalian genomes (25 orders), including 30 species of Cetacea. Our findings indicate that cetaceans and other mammals with atypical biorhythms, thermal constraints and/or energy metabolism, have accumulated deleterious mutations in CORT. In light of the pleiotropic action of this neuropeptide, we suggest that this inactivation contributed to a plethora of phenotypic adjustments to accommodate adaptive solutions to specific ecological niches., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Fisheries bycatch has been identified as the greatest threat to marine mammals worldwide. Characterizing the impacts of bycatch on marine mammals is challenging because it is difficult to both observe and quantify, particularly in small-scale fisheries where data on fishing effort and marine mammal abundance and distribution are often limited. The lack of risk frameworks that can integrate and visualize existing data have hindered the ability to describe and quantify bycatch risk. Here, we describe the design of a new geographic information systems tool built specifically for the analysis of bycatch in small-scale fisheries, called Bycatch Risk Assessment (ByRA). Using marine mammals in Malaysia and Vietnam as a test case, we applied ByRA to assess the risks posed to Irrawaddy dolphins (Orcaella brevirostris) and dugongs (Dugong dugon) by five small-scale fishing gear types (hook and line, nets, longlines, pots and traps, and trawls). ByRA leverages existing data on animal distributions, fisheries effort, and estimates of interaction rates by combining expert knowledge and spatial analyses of existing data to visualize and characterize bycatch risk. By identifying areas of bycatch concern while accounting for uncertainty using graphics, maps and summary tables, we demonstrate the importance of integrating available geospatial data in an accessible format that taps into local knowledge and can be corroborated by and communicated to stakeholders of data-limited fisheries. Our methodological approach aims to meet a critical need of fisheries managers: to identify emergent interaction patterns between fishing gears and marine mammals and support the development of management actions that can lead to sustainable fisheries and mitigate bycatch risk for species of conservation concern., Competing Interests: The authors have declared that no competing interests exist.

A wide range of anthropogenic structures exist in the marine environment with the extent of these set to increase as the global offshore renewable energy industry grows. Many of these pose acute risks to marine wildlife; for example, tidal energy generators have the potential to injure or kill seals and small cetaceans through collisions with moving turbine parts. Information on fine scale behaviour of animals close to operational turbines is required to understand the likely impact of these new technologies. There are inherent challenges associated with measuring the underwater movements of marine animals which have, so far, limited data collection. Here, we describe the development and application of a system for monitoring the three-dimensional movements of cetaceans in the immediate vicinity of a subsea structure. The system comprises twelve hydrophones and software for the detection and localisation of vocal marine mammals. We present data demonstrating the systems practical performance during a deployment on an operational tidal turbine between October 2017 and October 2019. Three-dimensional locations of cetaceans were derived from the passive acoustic data using time of arrival differences on each hydrophone. Localisation accuracy was assessed with an artificial sound source at known locations and a refined method of error estimation is presented. Calibration trials show that the system can accurately localise sounds to 2m accuracy within 20m of the turbine but that localisations become highly inaccurate at distances greater than 35m. The system is currently being used to provide data on rates of encounters between cetaceans and the turbine and to provide high resolution tracking data for animals close to the turbine. These data can be used to inform stakeholders and regulators on the likely impact of tidal turbines on cetaceans., Competing Interests: The authors have declared that no competing interests exist.

The cetaceans display a wide variety of lifestyles, especially with regard to their feeding behavior. However, the evolutionary process of the feeding behavior in cetaceans is still poorly understood, in part because reconstructing the feeding behavior of extinct taxa remains difficult. In cetaceans, cranium mobility relative to the trunk largely depends on the range of motion permitted by the atlanto-occipital joint, given the lack of flexibility of the cervical series. In this study, we examined 56 extant cetacean skeletal specimens from 30 species in 25 genera and nine families in order to investigate the relationships between anatomical traits and feeding behavior. Our results suggest that the range of dorso-ventral motion allowed by the atlanto-occipital joint (ROM) depends on prey habitat and the feeding technique of cetaceans. Cetaceans feeding on benthic/demersal prey had a relatively large ROM compared with those feeding on pelagic prey. In addition, ROM was largest in raptorial feeders, intermediate in suction feeders, and smallest in ram-filter feeders. Among raptorial feeders, ROM tended to be larger in taxa that facultatively tear off the prey's flesh compared with taxa that swallow their prey whole. Therefore, we conclude that ROM is a powerful tool to reliably reconstruct the feeding behavior of extinct cetacean taxa., (© 2019 Anatomical Society.)

Displays of maximum swimming speeds are rare in the laboratory and the wild, limiting our understanding of the top-end athletic capacities of aquatic vertebrates. However, jumps out of the water - exhibited by a diversity of fish and cetaceans - might sometimes represent a behaviour comprising maximum burst effort. We collected data on such breaching behaviour for 14 fish and cetacean species primarily from online videos, to calculate breaching speed. From newly derived formulae based on the drag coefficient and hydrodynamic efficiency, we also calculated the associated power. The fastest breaching speeds were exhibited by species 2 m in length, peaking at nearly 11 m s -1 ; as species size decreases below this, the fastest breaches become slower, while species larger than 2 m do not show a systematic pattern. The power associated with the fastest breaches was consistently ∼50 W kg -1 (equivalent to 200 W kg -1 muscle) in species from 20 cm to 2 m in length, suggesting that this value may represent a universal (conservative) upper boundary. And it is similar to the maximum recorded power output per muscle mass recorded in any species of similar size, suggesting that some breaches could indeed be representative of maximum capability., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Passive acoustic monitoring has become an important data collection method, yielding massive datasets replete with biological, environmental and anthropogenic information. Automated signal detectors and classifiers are needed to identify events within these datasets, such as the presence of species-specific sounds or anthropogenic noise. These automated methods, however, are rarely a complete substitute for expert analyst review. The ability to visualize and annotate acoustic events efficiently can enhance scientific insights from large, previously intractable datasets. A MATLAB-based graphical user interface, called DetEdit, was developed to accelerate the editing and annotating of automated detections from extensive acoustic datasets. This tool is highly-configurable and multipurpose, with uses ranging from annotation and classification of individual signals or signal-clusters and evaluation of signal properties, to identification of false detections and false positive rate estimation. DetEdit allows users to step through acoustic events, displaying a range of signal features, including time series of received levels, long-term spectral averages, time intervals between detections, and scatter plots of peak frequency, RMS, and peak-to-peak received levels. Additionally, it displays either individual, or averaged sound pressure waveforms, and power spectra within each acoustic event. These views simultaneously provide analysts with signal-level detail and encounter-level context. DetEdit creates datasets of signal labels for further analyses, such as training classifiers and quantifying occurrence, abundances, or trends. Although designed for evaluating underwater-recorded odontocete echolocation click detections, DetEdit can be adapted to almost any stereotyped impulsive signal. Our software package complements available tools for the bioacoustic community and is provided open source at https://github.com/MarineBioAcousticsRC/DetEdit., Competing Interests: The authors have declared that no competing interests exist

Cetaceans possess brains that rank among the largest to have ever evolved, either in terms of absolute mass or relative to body size. Cetaceans have evolved these huge brains under relatively unique environmental conditions, making them a fascinating case study to investigate the constraints and selection pressures that shape how brains evolve. Indeed, cetaceans have some unusual neuroanatomical features, including a thin but highly folded cerebrum with low cortical neuron density, as well as many structural adaptations associated with acoustic communication. Previous reports also suggest that at least some cetaceans have an expanded cerebellum, a brain structure with wide-ranging functions in adaptive filtering of sensory information, the control of motor actions, and cognition. Here, we report that, relative to the size of the rest of the brain, both the cerebrum and cerebellum are dramatically enlarged in cetaceans and show evidence of co-evolution, a pattern of brain evolution that is convergent with primates. However, we also highlight several branches where cortico-cerebellar co-evolution may be partially decoupled, suggesting these structures can respond to independent selection pressures. Across cetaceans, we find no evidence of a simple linear relationship between either cerebrum and cerebellum size and the complexity of social ecology or acoustic communication, but do find evidence that their expansion may be associated with dietary breadth. In addition, our results suggest that major increases in both cerebrum and cerebellum size occurred early in cetacean evolution, prior to the origin of the major extant clades, and predate the evolution of echolocation., (© 2019 The Authors. Journal of Evolutionary Biology published by John Wiley & Sons Ltd on behalf of European Society for Evolutionary Biology.)

Cetacean stranding records can provide vital information on species richness and diversity through space and time. Here we collate stranding records from Victoria, Australia and assess them for temporal, spatial and demographic trends. Between 1920 and 2016, 424 stranding events involving 907 individuals were recorded across 31 Cetacea species from seven families, including five new species records for the state. Seven of these events were mass strandings, and six mother and calf strandings were recorded. Importantly, 48% of the species recorded are recognised as data deficient on the IUCN Red List. The most commonly recorded taxa were Tursiops spp. (n = 146) and Delphinus delphis (common dolphins, n = 81), with the greatest taxonomic richness (n = 24) and highest incidence of stranding events documented within the Otways mesoscale bioregion. We found no seasonal stranding patterns anywhere in the state. While our findings improve understanding of the spatial and temporal patterns of cetacean diversity within Victoria, we suggest greater effort to collect demographic data at stranding events in order to better study state-wide patterns through time. We conclude with guidelines for minimum data collection standards for future strandings to maximise information capture from each event., Competing Interests: The authors have declared that no competing interests exist.

Most cetacean species are wide-ranging and highly mobile, creating significant challenges for researchers by limiting the scope of data that can be collected and leaving large areas un-surveyed. Aerial surveys have proven an effective way to locate and study cetacean movements but are costly and limited in spatial extent. Here we present a semi-automated pipeline for whale detection from very high-resolution (sub-meter) satellite imagery that makes use of a convolutional neural network (CNN). We trained ResNet, and DenseNet CNNs using down-scaled aerial imagery and tested each model on 31 cm-resolution imagery obtained from the WorldView-3 sensor. Satellite imagery was tiled and the trained algorithms were used to classify whether or not a tile was likely to contain a whale. Our best model correctly classified 100% of tiles with whales, and 94% of tiles containing only water. All model architectures performed well, with learning rate controlling performance more than architecture. While the resolution of commercially-available satellite imagery continues to make whale identification a challenging problem, our approach provides the means to efficiently eliminate areas without whales and, in doing so, greatly accelerates ocean surveys for large cetaceans., Competing Interests: I have read the journal's policy and the authors of this manuscript have the following competing interests: GH, GN, CH, and VK work for companies which may consider a commercial application for this research in the future. This does not alter our adherence to PLOS ONE policies on sharing data and materials.

Cetaceans are fully aquatic predatory mammals that have successfully colonized virtually all marine habitats. Their adaptation to these habitats, so radically different from those of their terrestrial ancestors, can give us comparative insights into the evolution of female roles and kinship in mammalian societies. We provide a review of the diversity of such roles across the Cetacea, which are unified by some key and apparently invariable life-history features. Mothers are uniparous, while paternal care is completely absent as far as we currently know. Maternal input is extensive, lasting months to many years. Hence, female reproductive rates are low, every cetacean calf is a significant investment, and offspring care is central to female fitness. Here strategies diverge, especially between toothed and baleen whales, in terms of mother-calf association and related social structures, which range from ephemeral grouping patterns to stable, multi-level, societies in which social groups are strongly organized around female kinship. Some species exhibit social and/or spatial philopatry in both sexes, a rare phenomenon in vertebrates. Communal care can be vital, especially among deep-diving species, and can be supported by female kinship. Female-based sociality, in its diverse forms, is therefore a prevailing feature of cetacean societies. Beyond the key role in offspring survival, it provides the substrate for significant vertical and horizontal cultural transmission, as well as the only definitive non-human examples of menopause. This article is part of the theme issue 'The evolution of female-biased kinship in humans and other mammals'.

Dorso-ventral oscillations of cetacean caudal flukes generate lift-based thrust for swimming. Movements of the flukes are actuated by epaxial and hypaxial muscles through caudal tendons inserting onto vertebrate in the peduncle. To determine if the caudal tendons in the peduncle affect the flexibility of the flukes, we must understand how the tendons from axial muscles insert onto the caudal vertebrate. The purpose of this study was to provide a detailed description of the various tendons within the cetacean peduncle with regard to their role in swimming and flexibility. Dissection of the peduncle and flukes of multiple odontocete species showed that there were two distinct epaxial tendon sets within the peduncle: (1) extensor caudae medialis tendon (ECM) and (2) extensor caudae lateralis tendon (ECL). There is one distinct hypaxial tendon set, the medial tendon of the hypaxialis lumborum (MHL). The ECM and MHL tendons inserted serially onto caudal vertebrae while the ECL inserted exclusively onto the terminal vertebrae posterior to the fluke insertion. It is typical that tendons insert onto bone, however, the connection to the core fibrous layer of the flukes suggests an element of active control of the flexibility of the flukes via the axial muscles. Tension from muscular contraction transmitted through the tendons could affect both spanwise and chordwise flexibility. Changing flexibility could modulate thrust and efficiency over an extended operation range of swimming speeds in cetaceans., (© 2019 Wiley Periodicals, Inc.)

Active acoustic sensors are widely used in oceanographic and environmental studies. Although many have nominal operating frequencies above the range of marine mammal hearing, they can produce out-of-band sound that may be audible to marine mammals. Acoustic emissions from four active acoustic transducers were characterized and compared to marine mammal hearing thresholds. All four transducers had nominal operating frequencies above the reported upper limit of marine mammal hearing, but produced measurable sound below 160 kHz. A spatial map of the acoustic emissions of each sonar is used to evaluate potential effects on marine mammal hearing when the transducer is continuously operated from a stationary platform. Based on the cumulative sound exposure level metric, the acoustic emissions from the transducers are unlikely to cause temporary threshold shifts in marine mammals, but could affect animal behavior. The extent of audibility is estimated to be, at most, on the order of 100 m., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Hydrofoil-shaped limbs (flipper-hydrofoils) have evolved independently several times in secondarily marine tetrapods and generally fall into two functional categories: (1) those that produce the majority of thrust during locomotion (propulsive flipper-hydrofoils); (2) those used primarily to steer and resist destabilizing movements such as yaw, pitch, and roll (controller flipper-hydrofoils). The morphological differences between these two types have been poorly understood. Theoretical and experimental studies on engineered hydrofoils suggest that flapping hydrofoils with a flexible trailing edge are more efficient at producing thrust whereas hydrofoils used in steering and stabilization benefit from a more rigid one. To investigate whether the trailing edge is generally more flexible in propulsive flipper-hydrofoils, we compared the bone distribution along the chord in both flipper types. The propulsive flipper-hydrofoil group consists of the forelimbs of Chelonioidea, Spheniscidae, and Otariidae. The controller flipper-hydrofoil group consists of the forelimbs of Cetacea. We quantified bone distribution from radiographs of species representing more than 50% of all extant genera for each clade. Our results show that the proportion of bone in both groups is similar along the leading edge (0-40% of the chord) but is significantly less along the trailing edge for propulsive flipper-hydrofoils (40-80% of the chord). Both flipper-hydrofoil types have little to no bony tissue along the very edge of the trailing edge (80-100% of the chord). This suggests a relatively flexible trailing edge for propulsive flipper-hydrofoils compared to controller flipper-hydrofoils in line with findings from prior studies. This study presents a morphological correlate for inferring flipper-hydrofoil function in extinct taxa and highlights the importance of a flexible trailing edge in the evolution of propulsive flipper-hydrofoils in marine tetrapods., (© 2019 Wiley Periodicals, Inc.)

Very little is known about marine mammal susceptibility to primary blast injury (PBI) except in rare cases of opportunistic studies. As a result, traditional analysis techniques relied on methods developed more than 30 years ago using terrestrial mammals as surrogates. Modeling tools available today have the computing power to vastly improve calculation of safe ranges and injury zones from underwater explosions (UNDEX) employing morphologically accurate proxies with material properties similar to marine mammal tissues. The Dynamic System Mechanics Advanced Simulation (DYSMAS) fluid-structure interaction (FSI) software is being used to simulate the complex phenomena of UNDEX, shock wave, and bubble pulse propagation through the water and transmission of energy to a cetacean focusing on the dynamic response of the thoracic cavity and air-filled lungs to a shock wave. The approach integrates fluid and structural analyses with the material properties of blubber, bone, and muscle using marine mammal morphometrics to eliminate unnecessary assumptions made during more traditional approaches to analysis developed before these types of data and computational power were available. DYSMAS analyses of a 1D gas bubble surrounded by water was found to closely match the classical bubble dynamics models. Further, DYSMAS models of a spherical gas bubble surrounded by tissue and rib structure demonstrate a global radial oscillation of the gas bubble, but also show significant local deflection and material strain in response to the UNDEX loading. The intended result of the investigation is an improved and scientifically defensible understanding of the effects of UNDEX on marine mammals. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc. Anat Rec, 302:718-734, 2019. Published 2018. This article is a U.S. Government work and is in the public domain in the USA., (Published 2018. This article is a U.S. Government work and is in the public domain in the USA.)

Marine litter is a pollution problem affecting thousands of marine species in all the world's seas and oceans. Marine litter, in particular plastic, has negative impacts on marine wildlife primarily due to ingestion and entanglement. Since most marine mammal species negatively interact with marine litter, a first workshop under the framework of the European Cetacean Society Conference, was held in 2017 to bring together the main experts on the topic of marine mammals and marine litter from academic and research institutes, non-governmental organisations, foundations and International Agreements. The workshop was devoted to defining the impact of marine litter on marine mammals by reviewing current knowledge, methodological advances and new data available on this emerging issue. Some case studies were also presented from European waters, such as seals and cetaceans in the North, Baltic, and Mediterranean Seas. Here, we report the main findings of the workshop, including a discussion on the research needs, the main methodological gaps, an overview of new techniques for detecting the effects of marine litter (including microplastics) on marine mammals and, also, the use of citizen science to drive awareness. The final recommendations aim to establish priority research, to define harmonised methods to detect marine litter and microplastics, enforce networking among institutions and support data sharing. The information gathered will enhance awareness and communication between scientists, young people, citizens, other stakeholders and policy makers, and thereby facilitate better implementation of international directives (e.g., the Marine Strategy Framework Directive) in order to answer the question about the actual status of our oceans and finding solutions., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Studying ontogeny in both extant and extinct species can unravel the mechanisms underlying mammal diversification and specialization. Among mammalian clades, Cetartiodactyla encompass species with a wide range of adaptations, and ontogenetic evidence could clarify longstanding debates on the origins of modern specialized families. Here, we study the evolution of dental eruption patterns in early diverging cetartiodactyls to assess the ecological and biological significance of this character and shed new light on phylogenetic issues. After investigation of the ontogenetic dental series of 63 extinct genera, our parsimony reconstructions of eruption state evolution suggest that the eruption of molars before permanent premolars represents a plesiomorphic condition within Cetartiodactyla. This result substantially differs from a previous study based on modern species only. As a result, the presence of this pattern in most ruminants might represent an ancestral condition contributing to their specialized herbivory, rather than an original adaptation. In contrast, the late eruption of molars in hippopotamoids is more likely related to biological aspects, such as increases in body mass and slower pace of life. Our study mainly shows that eruption sequences reliably characterize higher level cetartiodactyl taxa and could represent a new source of phylogenetic characters, especially to disentangle the origin of hippopotamoids and cetaceans.

: Melatonin, the hormone of darkness, is a peculiar molecule found in most living organisms. Emerging as a potent broad-spectrum antioxidant, melatonin was repurposed into extra roles such as the modulation of circadian and seasonal rhythmicity, affecting numerous aspects of physiology and behaviour, including sleep entrainment and locomotor activity. Interestingly, the pineal gland-the melatonin synthesising organ in vertebrates-was suggested to be absent or rudimentary in some mammalian lineages, including Cetacea. In Cetacea, pineal regression is paralleled by their unique bio-rhythmicity, as illustrated by the unihemispheric sleeping behaviour and long-term vigilance. Here, we examined the genes responsible for melatonin synthesis ( Aanat and Asmt ) and signalling ( Mtnr1a and Mtnr1b ) in 12 toothed and baleen whale genomes. Based on an ample genomic comparison, we deduce that melatonin-related gene modules are eroded in Cetacea., Competing Interests: The authors declare no conflict of interest.

In this work, a convolutional neural network based method is proposed to automatically detect odontocetes echolocation clicks by analyzing acoustic data recordings from a passive acoustic monitoring system. The neural network was trained to distinguish between click and non-click clips and was subsequently converted to a full-convolutional network. The performance of the proposed network was evaluated using synthetic data and real audio recordings. The experimental results indicate that the proposed method works stably with echolocation clicks of different species.

In mammalian evolutionary history, Cetacea (whales, dolphins and porpoises) achieved astonishing success by adapting to an aquatic environment. One unique characteristic of cetaceans, contributing to this adaptive success, is efficient lipid utilization. Here, we report a comparative genetic analysis of five aquatic and five terrestrial Cetartiodactyla species using 144 genes associated with lipid metabolism. Mutation ratio (d N /d S ), amino acid substitution in functional domains and metabolic pathways were evaluated using branch-site model in PAML, Pfam and KEGG, respectively. Our tests detected 20 positively selected genes in Cetacea compared to 11 in Bovidae with little overlap between the lineages. We identified lineage-specific patterns of amino acid substitutions and functional domains that were mutually exclusive between cetaceans and bovids, supporting divergent evolution of lipid metabolism since the divergence of these taxa from a common ancestor. Moreover, a pathway analysis showed that the identified genes in cetaceans were associated with lipid digestion, lipid storage and energy-producing pathways. This study emphasizes the evolutionary context of lipid metabolism modification of cetaceans and provides a foundation for future studies of elucidating the adapted biological mechanisms of cetacean lipid metabolism and a framework for incorporating ecological context into studies aimed at investigating adaptive evolution., (© 2018 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2018 European Society For Evolutionary Biology.)

Global warming is significantly altering arctic marine ecosystems. Specifically, the precipitous loss of sea ice is creating a dichotomy between ice-dependent polar bears and pinnipeds that are losing habitat and some cetaceans that are gaining habitat. While final outcomes are hard to predict for the many and varied marine mammal populations that rely on arctic habitats, we suggest a simplified framework to assess status, based upon ranking a population's size, range, behavior, and health. This basic approach is proposed as a means to prioritize and expedite conservation and management efforts in an era of rapid ecosystem alteration., Competing Interests: The authors have declared that no competing interests exist.

Some aquatic mammals appear to care for their dead, whereas others abandon their live offspring when conditions are unfavourable. This incredible variety in behaviours suggests the importance of comparing and contrasting mechanisms driving death-related behaviours among these species. We reviewed 106 cases of aquatic mammals (81 cetaceans and 25 non-cetaceans) reacting to a death event, and extrapolated 'participant' ( age class , sex , relationship and decomposition ) and 'social' characteristics ( escorting , calf dependence , alloparental care , herding and dispersal patterns ) from published and unpublished literature. A multiple correspondence analysis (MCA) was performed to explore the relationships between these characteristics and death-related behaviours, with species clustered based on MCA scores. Results showed that both cetaceans and non-cetaceans react to death but in different ways. Non-cetaceans, characterized by a short maternal investment, were observed to protect the dead (defending it from external attacks), while cetaceans spent much longer with their offspring and display carrying (hauling, spinning, mouthing with the carcass and diving with it) and breathing-related (lifting and sinking the carcass) activities with the dead generally in association with other conspecifics. Our work emphasizes the need of increased documentation of death-related cases around the world to improve our understanding of aquatic mammals and their responses to death.This article is part of the theme issue 'Evolutionary thanatology: impacts of the dead on the living in humans and other animals'., (© 2018 The Author(s).)

We used ecosystem sampling during systematic surveys and opportunistic focal follows, comparison tests, and random forest models to evaluate fin whale (Balaenoptera physalus) and humpback whale (Megaptera novaeangliae) habitat associations within an inland feeding ground (Kitimat Fjord System, British Columbia, Canada). Though these species are sympatric and share a common prey source, they were attuned to different aspects of the local habitat. The fin whales were associated with habitat properties reminiscent of the open ocean. Humpback whales, in contrast, were associated with features more commonly associated with the inland waters of fjords. Fixed habitat features, such as seafloor depth and distance from the fjord mouth, were the most important predictors of fin whale presence, but fixed and dynamic variables, such as surface properties, predicted humpback whale presence with equal (moderate) success. With the exception of strong salinity gradients for humpback whales, habitat conditions were poor predictors of feeding state. Fin whales practiced a spatially confined, seasonally stable, and thus more predictable use of certain channels within the fjord system. These findings are compatible with site loyal behavior, which is interesting in light of the species' historical, unique use of this fjord system. The relatively lackluster performance of humpback-habitat models, coupled with the importance of oceanographic properties, makes the humpback's habitat use strategy more uncertain. The fact that two sympatric species sharing a common prey source exhibited different habitat use strategies suggests that at least one species was informed by something in addition to prey. Given that the two species are attuned to different aspects of the fjord habitat, their responses to habitat changes, including anthropogenic impacts, would likely be different in both nature and degree. Our findings highlight the value of comparative studies and the complexity of rorqual habitat use, which must be understood in order for critical habitat to be identified and protected., (Copyright © 2018. Published by Elsevier Ltd.)

For about 119 species of cetaceans and other aquatic animals, sound is the key source of learning about the environment, navigation, communication, foraging, and avoiding predators. However, in the recent era, the introduction of large quantities of anthropogenic noise into the ocean has significantly altered the ocean's acoustic environment. The anthropogenic noises travel very long distances, blanketing enormous areas. This can affect cetaceans, either by direct killing or compromising hearing, navigation, communication, predation, as well as normal behaviors. It has been suggested that acoustic pollution could possibly negatively affect cetacean reproduction, which is harmful for endangered and threatened species. However, it is still unknown how acoustic pollution can suppress cetacean reproduction. This is the first comprehensive review article, which focuses on the possible consequences affecting the reproduction of marine mammals resulting from acoustic pollution.