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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’.

Narwhals (Monodon monoceros) are deep-diving Arctic cetaceans that migrate seasonally between summering and wintering grounds. The Baffin Bay population overwinters in southern Baffin Bay and Davis Strait, where they are known to forage on high-energy benthic prey. Studying narwhal winter behaviour and prey preference has been challenged by their remote distribution and limited lifespan of satellite tags deployed in summer, restricting data on their habitat use and foraging strategies. Since prey consumption is thought to peak in the winter, understanding narwhal diet plasticity in a rapidly changing environment like Baffin Bay is critical. This study developed unique methods to examine four years of irregular satellite telemetry data from 22 narwhals tagged in their summering grounds. Locations and recorded diving data from the overwintering area were isolated, and a hidden Markov model was used to define three behaviours (\"surface\", \"pelagic\", and \"deep-water\" diving). We further examined the effects of five covariates on these behaviours to provide insight into the spatial patterns of narwhal winter prey preference. Narwhal behaviours were dominated by diving, with 37% of their time spent in pelagic waters and 40% in deep-water, while only 22% of their time was spent in surface related behaviours. Deep-water behaviours increased later in the day and into the winter season and occurred frequently in the center trough of Baffin Bay before (66°- 69°) and across Davis Strait (65° - 67°). In contrast, pelagic behaviours declined as the winter season progressed and occurred earlier in the day. Narwhals occupying the northern overwintering area exhibited more pelagic behaviours, despite it being deeper, suggesting different foraging strategies across their winter range. Our study identified behaviours suggestive of a variable winter diet and provided insight on the spatial nature of these behaviours across the winter season. The methods developed in this study present new opportunities for analysing lower resolution satellite tracking data. With advancements in bio-logging technology and remote field methods, the ability to successfully document changes in winter space use and fine-scale foraging behaviours may be possible for narwhal in the future.

As human activities impact virtually every animal habitat on the planet, identifying species at-risk from disturbance is a priority. Cetaceans are an example taxon where responsiveness to anthropogenic noise can be severe but highly species and context specific, with source–receiver characteristics such as hearing sensitivity only partially explaining this variability. Here, we predicted that ecoevolutionary factors that increase species responsiveness to predation risk also increase responsiveness to anthropogenic noise. We found that reductions in intense-foraging time during exposure to 1- to 4-kHz naval sonar and predatory killer whale sounds were highly correlated (r = 0.92) across four cetacean species. Northern bottlenose whales ceased foraging completely during killer whale and sonar exposures, followed by humpback, long-finned pilot, and sperm whales, which reduced intense foraging by 48 to 97%. Individual responses to sonar were partly predicted by species-level responses to killer whale playbacks, implying a similar level of perceived risk. The correlation cannot be solely explained by hearing sensitivity, indicating that species- and context-specific antipredator adaptations also shape cetacean responses to human-made noise. Species that are more responsive to predator presence are predicted to be more disturbance sensitive, implying a looming double whammy for Arctic cetaceans facing increased anthropogenic and predator activity with reduced ice cover.

The remarkable fossil record of whales and dolphins (Cetacea) has made them an exemplar of macroevolution. Although their overall adaptive transition from terrestrial to fully aquatic organisms is well known, this is not true for the radiation of modern whales. Here, we explore the diversification of extant cetaceans by constructing a robust molecular phylogeny that includes 87 of 89 extant species. The phylogeny and divergence times are derived from nuclear and mitochondrial markers, calibrated with fossils. We find that the toothed whales are monophyletic, suggesting that echolocation evolved only once early in that lineage some 36–34 Ma. The rorqual family (Balaenopteridae) is restored with the exclusion of the gray whale, suggesting that gulp feeding evolved 18–16 Ma. Delphinida, comprising all living dolphins and porpoises other than the Ganges/Indus dolphins, originated about 26 Ma; it contains the taxonomically rich delphinids, which began diversifying less than 11 Ma. We tested 2 hypothesized drivers of the extant cetacean radiation by assessing the tempo of lineage accumulation through time. We find no support for a rapid burst of speciation early in the history of extant whales, contrasting with expectations of an adaptive radiation model. However, we do find support for increased diversification rates during periods of pronounced physical restructuring of the oceans. The results imply that paleogeographic and paleoceanographic changes, such as closure of major seaways, have influenced the dynamics of radiation in extant cetaceans.

To satisfy the needs of sleeping underwater, marine mammals, including cetaceans, sirenians, and pinnipeds, have evolved an unusual form of sleep, known as unihemispheric slow-wave sleep (USWS), in which one brain hemisphere is asleep while the other is awake. All aquatic cetaceans have only evolved USWS without rapid eye movement (REM) sleep, whereas aquatic sirenians and amphibious pinnipeds display both bihemispheric slow-wave sleep (BSWS) and USWS, as well as REM sleep. However, the molecular genetic changes underlying USWS remain unknown. The present study investigated the evolution of eight canonical circadian genes and found that positive selection occurred mainly within cetacean lineages. Furthermore, convergent evolution was observed in lineages with USWS at three circadian clock genes. Remarkably, in vitro assays showed that cetacean-specific mutations increased the nuclear localization of zebrafish clocka , and enhanced the transcriptional activation activity of Clocka and Bmal1a. In vivo , transcriptome analysis showed that the overexpression of the cetacean-specific mutant clocka ( clocka -mut) caused the upregulation of the wakefulness-promoting glutamatergic genes and the differential expression of multiple genes associated with sleep regulation. In contrast, the GABAergic and cholinergic pathways, which play important roles in promoting sleep, were downregulated in the bmal1a -mut-overexpressing zebrafish. Concordantly, sleep time of zebrafish overexpressing clocka -mut and bmal1a -mut were significantly less than the zebrafish overexpressing the wild-type genes, respectively. These findings support our hypothesis that canonical circadian clock genes may have evolved adaptively to enhance circadian regulation ability relating to sleep in cetaceans and, in turn, contribute to the formation of USWS.

Male genitalia evolve rapidly, probably as a result of sexual selection. Whether this pattern extends to the internal infrastructure that influences genital movements remains unknown. Cetaceans (whales and dolphins) offer a unique opportunity to test this hypothesis: since evolving from land-dwelling ancestors, they lost external hind limbs and evolved a highly reduced pelvis that seems to serve no other function except to anchor muscles that maneuver the penis. Here, we create a novel morphometric pipeline to analyze the size and shape evolution of pelvic bones from 130 individuals (29 species) in the context of inferred mating system. We present two main findings: (1) males from species with relatively intense sexual selection (inferred by relative testes size) tend to evolve larger penises and pelvic bones compared to their body length, and (2) pelvic bone shape has diverged more in species pairs that have diverged in inferred mating system. Neither pattern was observed in the anterior-most pair of vertebral ribs, which served as a negative control. This study provides evidence that sexual selection can affect internal anatomy that controls male genitalia. These important functions may explain why cetacean pelvic bones have not been lost through evolutionary time.

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.

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.