‘Birds’ of two feathers: Avicranium renestoi and the paraphyly of bird-headed reptiles (Diapsida: ‘Avicephala’)

The anatomy of Late Triassic drepanosauromorphs is re-examined, with a focus on the previously published surface models of the holotype of Avicranium renestoi from the Norian of North America. We comment on the cranial anatomy of this taxon and propose a new reconstruction of the skull and mandible. Contrary to previous interpretations, the entire rostrum and most of the palate are not preserved in this specimen. We also suggest that some proposed plesiomorphic characters may result from incomplete ossification due to immaturity. These new observations are compiled into a new morphological phylogenetic dataset designed to address the monophyly of ‘Avicephala’, the group comprising the Late Permian gliding reptiles Weigeltisauridae, and the Late Triassic chameleon-like Drepanosauromorpha. We recover Weigeltisauridae as stem-saurian diapsids and Drepanosauromorpha as sister-group to Trilophosauridae among archosauromorphs, thus implying the paraphyly of ‘Avicephala’. Drepanosauromorphs and trilophosaurids are recovered as sister-taxa for the first time, as supported by several cranial and postcranial synapomorphies. This new phylogenetic position of Drepanosauromorpha reduces the group’s ghost lineage that now does not necessarily cross the Permian–Triassic boundary. However, much remains unknown of the early history of trilophosaurids and drepanosauromorphs, and of the evolution of arboreality in Triassic archosauromorph reptiles.