Covariation in the Craniocervical Junction of Carnivora

The craniocervical junction is the transition between the skull and the vertebral column that provides mobility while maintaining sufficient stability (i.e., protection of the brainstem and the spinal cord). The key elements involved are the occiput, the first cervical vertebra (CV1, atlas) and the second cervical vertebra (CV2, axis). The two vertebrae forming the atlas-axis complex are distinct in their morphology and differences in form have been linked to differences in ecological function in mammals. Here, we quantified the morphological diversity of the cranium, CV1 and CV2 in a sample of Carnivora using 3D geometric morphometrics to reveal phylogenetic and ecological patterns. Our results indicate that the observed variation in CV2 is related to the taxonomic diversity (i.e., strong phylogenetic signal), whereas variation in CV1 appears to be decoupled from species diversity in Carnivora and, thus, is likely to reflect a functional signal. The phylogenetically informed correlation analyses showed an association between the CV1 morphology and diet. Taxa that primarily feed on large prey tend to have larger transverse processes on CV1 which provides larger muscle attachment areas and may correlate with stronger muscles. The latter needs to be verified by future quantitative covariation analyses between bone and muscle data. Morphological peculiarities within Pinnipedia and Mustelidae could be explained by differences in terrestrial locomotion between Phocidae and Otariidae and the exceptional defensive behavior (i.e., handstanding) in Mephitidae. Despite differences in the degree of morphological diversity, covariation between cranium, CV1 and CV2 morphology is consistently high (≥ 0.82) highlighting that overall, the craniocervical junction is an integrated structure, but there are traits that are not constrained.