The evolution and role of the hyposphene-hypantrum articulation in Archosauria: phylogeny, size and/or mechanics?

Abstract

Living members of Archosauria, the reptile clade containing Crocodylia and Aves, have a wide range of skeletal morphologies, ecologies and body size. The range of body size greatly increases when extinct archosaurs are included, because extinct Archosauria includes the largest members of any terrestrial vertebrate group (e.g. 70-tonne titanosaurs, 20-tonne theropods). Archosaurs evolved various skeletal adaptations for large body size, but these adaptations varied among clades and did not always appear consistently with body size or ecology. Modification of intervertebral articulations, specifically the presence of a hyposphene-hypantrum articulation between trunk vertebrae, occurs in a variety of extinct archosaurs (e.g. non-avian dinosaurs, pseudosuchians). We surveyed the phylogenetic distribution of the hyposphene-hypantrum to test its relationship with body size. We found convergent evolution among large-bodied clades, except when the clade evolved an alternative mechanism for vertebral bracing. For example, some extinct lineages that lack the hyposphene-hypantrum articulation (e.g. ornithischians) have ossified tendons that braced their vertebral column. Ossified tendons are present even in small taxa and in small-bodied juveniles, but large-bodied taxa with ossified tendons reached those body sizes without evolving the hyposphene-hypantrum articulation. The hyposphene-hypantrum was permanently lost in extinct crownward members of both major archosaur lineages (i.e. Crocodylia and Aves) as they underwent phyletic size decrease, changes in vertebral morphology and shifts in ecology.