Vertebral morphology in extant porpoises: Radiation and functional implications.

Vertebral morphology has profound biomechanical implications and plays an important role in adaptation to different habitats and foraging strategies for cetaceans. Extant porpoise species (Phocoenidae) display analogous evolutionary patterns in both hemispheres associated with convergent evolution to coastal versus oceanic environments. We employed 3D geometric morphometrics to study vertebral morphology in five porpoise species with contrasting habitats: the coastal Indo-Pacific finless porpoise (Neophocaena phocaenoides); the mostly coastal harbor porpoise (Phocoena phocoena) and Burmeister's porpoise (Phocoena spinipinnis); and the oceanic spectacled porpoise (Phocoena dioptrica) and Dall's porpoise (Phocoenoides dalli). We evaluated the radiation of vertebral morphology, both in size and shape, using multivariate statistics. We supplemented data with samples of an early-radiating delphinoid species, the narwhal (Monodon monoceros); and an early-radiating delphinid species, the white-beaked dolphin (Lagenorhynchus albirostris). Principal component analyses were used to map shape variation onto phylogenies, and phylogenetic constraints were investigated through permutation tests. We established links between vertebral morphology and movement patterns through biomechanical inferences from morphological presentations. We evidenced divergence in size between species with contrasting habitats, with coastal species tending to decrease in size from their estimated ancestral state, and oceanic species tending to increase in size. Regarding vertebral shape, coastal species had longer centra and shorter neural processes, but longer transverse processes, while oceanic species tended to have disk-shaped vertebrae with longer neural processes. Within Phocoenidae, the absence of phylogenetic constraints in vertebral morphology suggests a high level of evolutionary lability. Overall, our results are in accordance with the hypothesis of speciation within the family from a coastal ancestor, through adaptation to particular habitats. Variation in vertebral morphology in this group of small odontocetes highlights the importance of environmental complexity and particular selective pressures for the speciation process through the development of adaptations that minimize energetic costs during locomotion and prey capture.

Head adaptation for sound production and feeding strategy in dolphins (Odontoceti: Delphinida).

Head morphology in toothed whales evolved under selective pressures on feeding strategy and sound production. The postnatal development of the skull (n = 207) and mandible (n = 219) of six Delphinida species which differ in feeding strategy but exhibit similar sound emission patterns, including two narrow-band high-frequency species, were investigated through 3D morphometrics. Morphological changes throughout ontogeny were demonstrated based on the main source of variation (i.e., prediction lines) and the common allometric component. Multivariate trajectory analysis with pairwise comparisons between all species was performed to evaluate specific differences on the postnatal development of skulls and mandibles. Changes in the rostrum formation contributed to the variation (skull: 49%; mandible: 90%) of the entire data set and might not only reflect the feeding strategy adopted by each lineage but also represents an adaptation for sound production and reception. As an important structure for directionality of sound emissions, this may increase directionality in raptorial feeders. Phylogenetic generalized least squares analyses indicated that shape of the anterior portion of the skull is strongly dependent on phylogeny and might not only reflect feeding mode, but also morphological adaptations for sound production, particularly in raptorial species. Thus, postnatal development seems to represent a crucial stage for biosonar maturation in some raptorial species such as Pontoporia blainvillei and Sousa plumbea. The ontogeny of their main tool for navigation and hunting might reflect their natural history peculiarities and thus potentially define their main vulnerabilities to anthropogenic changes in the environment.