The Evolutionary Relationship Between Arm Vertebrae Shape and Ecological Lifestyle in Brittle Stars (Echinodermata: Ophiuroidea)

Ophiuroidea are one of the most diverse classes among extant echinoderms, characterized by their flexible arms composed of a series of ossicles called vertebrae, articulating with each other proximally and distally. Their arms show a wide range of motion, important for feeding and locomotion, associated with their epizoic and non-epizoic lifestyles. It remains to be explored to what degree the phenotypic variation in these ossicles also reflects adaptations to these lifestyles, rather than only their phylogenetic affinity. In this study, we analyzed the 3D shape variation of six arm vertebrae from the middle and distal parts of an arm in 12 species, belonging to the intertidal, subtidal and bathyal zones and showing epizoic and non-epizoic behaviors. A PERMANOVA indicated a significant difference in ossicle morphology between species and between lifestyles. A principal component analysis showed that the morphology of epizoic ophiuroids is distinct from non-epizoic ones; which may reflect variation in arm function related to these different lifestyles. The Phylogenetic MANOVA and phylogenetic signal analysis showed that shape variation in the vertebral articulation seems to reflect ecological and functional adaptations, whereas phylogeny controls more the lateral morphology of the vertebrae. This suggests a convergent evolution through ecological adaptation to some degree, indicating that some of these characters may have limited taxonomic value.