The earliest-diverging avemetatarsalian: a new osteoderm-bearing taxon from the Triassic (?Earliest Late Triassic) of Madagascar and the composition of avemetatarsalian assemblages prior to the radiation of dinosaurs

Abstract

AbstractUnderstanding the evolution of the earliest avemetatarsalian (bird-line) archosaurs and inferring the morphology of the last common ancestor of Archosauria are hampered by a poor fossil record in critical temporal intervals. Here we describe an early-diverging avemetatarsalian from the ?Earliest Late Triassic (~235 Ma) ‘basal Isalo II’/Makay Formation of Madagascar, which helps bridge these gaps. This taxon, Mambachiton fiandohana gen. et sp. nov., is represented by well-preserved postcranial material and possibly a postfrontal bone. Features of the neck region include anteroposteriorly elongated vertebrae with laterally expanded dorsal ends of the neural spines with three pairs of osteoderms per cervical vertebra, lying dorsal to those vertebrae. Inclusion of Mambachiton in a phylogenetic analysis of archosauromorphs recovers it at the base of Avemetatarsalia, outside of the aphanosaur + ornithodiran clade. This new specimen indicates that osteoderms were present in the earliest avemetatarsalians, but were lost in more crownward lineages. The plesiomorphic morphology of the taxon also underscores the difficulty of identifying early avemetatarsalians from incomplete skeletons. This early-diverging avemetatarsalian occurring together with a lagerpetid and silesaurid in the ‘basal Isalo II’/Makay Formation of Madagascar documents the co-occurrence of multiple non-dinosaurian avemetatarsalian clades in Gondwana near the Middle–Late Triassic transition. Translated abstract (Malagasy and French) is provided in the Supplementary information.