Social relationships and death-related behaviour in aquatic mammals: a systematic review.

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characteristics (escorting, calf dependence, alloparental care, herding and dispersal patterns) from published and unpublished literature.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'.

Rewriting the history of an extinction-was a population of Steller's sea cows (Hydrodamalis gigas) at St Lawrence Island also driven to extinction?

The Kommandorskiye Islands population of Steller's sea cow (Hydrodamalis gigas) was extirpated ca 1768 CE. Until now, Steller's sea cow was thought to be restricted in historic times to Bering and Copper Islands, Russia, with other records in the last millennium from the western Aleutian Islands. However, Steller's sea cow bone has been obtained by the authors from St Lawrence Island, Alaska, which is significantly further north. Bone identity was verified using analysis of mitochondrial DNA. The nitrogen-15 (δ(15)N)/carbon-13 (δ(13)C) values for bone samples from St Lawrence Island were significantly (p ≤ 0.05) different from Bering Island samples, indicating a second population. Bone samples were dated to between 1030 and 1150 BP (approx. 800-920 CE). The samples date from close to the beginning of the mediaeval warm period, which could indicate that the population at St Lawrence Island was driven to extinction by climate change. A warming of the climate in the area may have changed the availability of kelp; alternatively or in addition, the animals may have been driven to extinction by the expansion of the Inuit from the Bering Strait region, possibly due to opening waterways, maybe following bowhead whales (Balaena mysticetus), or searching for iron and copper. This study provides evidence for a previously unknown population of sea cows in the North Pacific within the past 1000 years and a second Steller's sea cow extirpation event in recent history.

Breaking constraint: axial patterning in Trichechus (Mammalia: Sirenia).

Meristic variation is often limited in serially homologous systems with high internal differentiation and high developmental modularity. The mammalian neck, an extreme example, has a fixed (at seven) count of diversely specialized segments. Imposition of the mammalian cervical constraint has been tentatively linked to the origin of the diaphragm, which is muscularized by cells that migrate from cervical somites during development. With six cervical vertebrae, the genus Trichechus (manatee) has apparently broken this constraint, although the mechanism of constraint escape is unknown. Hypotheses for the developmental origin of Trichechus cervical morphology include cervical rib 7 repatterning, a primaxial/abaxial patterning shift, and local homeosis at the cervical/thoracic boundary. We tested predictions of these hypotheses by documenting vertebral morphology, axial ossification patterns, regionalization of the postcranial skeleton, and the relationship of thoracic ribs to sternal subunits in a large data set of fetal and adult Trichechus and Dugong specimens. These observations forced rejection of all three hypotheses. We propose alternatively that a global slowing of the rate of somitogenesis reduced somite count and disrupted alignment of Hox-generated anatomical markers relative to somite (and vertebral) boundaries throughout the Trichechus column. This hypothesis is consistent with observations of the full range of traditional cervical morphologies in the six cervical vertebrae, conserved postcranial proportions, and column-wide reduction in count relative to its sister taxon, Dugong. It also suggests that the origin of the mammalian cervical constraint lies in patterning, not in count, and that Trichechus and the tree sloths have broken the constraint using different developmental mechanisms.

Reproductive cycles of marine mammals.

Marine mammals conform to the general mammalian reproductive system centered on the hypothalamic-pituitary-gonadal axis. Most marine mammals are long-lived and of large body size with lesser reproductive rates than many other animals, a consequence of their interaction with the marine environment where the demands of acquiring resources from the ocean must be balanced with the need for bearing offspring in a suitable place for survival. The degree of spatial and temporal separation of these life history phases in many species is a key feature of their ecology. The reproductive physiology of pinnipeds, cetaceans, sirenians, sea otters and polar bears has been more thoroughly characterized for the more accessible species.

Adaptive features of aquatic mammals' eye.

The eye of aquatic mammals demonstrates several adaptations to both underwater and aerial vision. This study offers a review of eye anatomy in four groups of aquatic animals: cetaceans (toothed and baleen whales), pinnipeds (seals, sea lions, and walruses), sirenians (manatees and dugongs), and sea otters. Eye anatomy and optics, retinal laminar morphology, and topography of ganglion cell distribution are discussed with particular reference to aquatic specializations for underwater versus aerial vision. Aquatic mammals display emmetropia (i.e., refraction of light to focus on the retina) while submerged, and most have mechanisms to achieve emmetropia above water to counter the resulting aerial myopia. As underwater vision necessitates adjusting to wide variations in luminosity, iris muscle contractions create species-specific pupil shapes that regulate the amount of light entering the pupil and, in pinnipeds, work in conjunction with a reflective optic tapetum. The retina of aquatic mammals is similar to that of nocturnal terrestrial mammals in containing mainly rod photoreceptors and a minor number of cones (however, residual color vision may take place). A characteristic feature of the cetacean and pinniped retina is the large size of ganglion cells separated by wide intercellular spaces. Studies of topographic distribution of ganglion cells in the retina of cetaceans revealed two areas of ganglion cell concentration (the best-vision areas) located in the temporal and nasal quadrants; pinnipeds, sirenians, and sea otters have only one such area. In general, the visual system of marine mammals demonstrates a high degree of development and several specific features associated with adaptation for vision in both the aquatic and aerial environments.