Cranial asymmetry in odontocetes: a facilitator of sonic exploration?

Directional cranial asymmetry is an intriguing condition that has evolved in all odontocetes which has mostly been associated with sound production for echolocation. In this study, we investigated how cranial asymmetry varies across odontocete species both in terms of quality (i.e., shape), and quantity (magnitude of deviation from symmetry). We investigated 72 species across all ten families of Odontoceti using two-dimensional geometric morphometrics. The average asymmetric shape was largely consistent across odontocetes - the rostral tip, maxillae, antorbital notches and braincase, as well as the suture crest between the frontal and interparietal bones were displaced to the right, whereas the nasal septum and premaxillae showed leftward shifts, in concert with an enlargement of the right premaxilla and maxilla. A clear phylogenetic signal related to asymmetric shape variation was identified across odontocetes using squared-change parsimony. The magnitude of asymmetry was widely variable across Odontoceti, with greatest asymmetry in Kogiidae, Monodontidae and Globicephalinae, followed by Physeteridae, Platanistidae and Lipotidae, while the asymmetry was lowest in Lissodelphininae, Phocoenidae, Iniidae and Pontoporiidae. Ziphiidae presented a wide spectrum of asymmetry. Generalized linear models explaining magnitude of asymmetry found associations with click source level while accounting for cranial size. Using phylogenetic generalized least squares, we reconfirm that source level and centroid size significantly predict the level of cranial asymmetry, with more asymmetric marine taxa generally consisting of bigger species emitting higher output sonar signal, i.e. louder sounds. Both characteristics theoretically support foraging at depth, the former by allowing extended diving and the latter being adaptive for prey detection at longer distances. Thus, cranial asymmetry seems to be an evolutionary pathway that allows odontocetes to devote more space for sound-generating structures associated with echolocation and thus increases biosonar search range and foraging efficiency beyond simple phylogenetic scaling predictions.