Skull osteology of the holotype of the rhynchosaur Hyperodapedon sanjuanensis (Sill, 1970) from the Upper Triassic Ischigualasto Formation of Argentina.

Rhynchosaurs are bulky quadrupedal herbivores that achieved a cosmopolitan distribution during the Middle and Late Triassic. Rhynchosaurids are characterized by a pair of premaxillae modified into an edentulous beak that had a bone-to-bone occlusion with the tips of the dentaries, and a specialized masticatory apparatus composed of groove(s) on the maxilla and ridges(s) on the dentary. The Argentinian fossil record of rhynchosaurs is abundant, but only two nominal species have been named so far. One of them, the hyperodapedontine Hyperodapedon sanjuanensis, is the most abundant tetrapod of the Ischigualasto Formation of northwestern Argentina. However, the anatomy of H. sanjuanensis remains poorly studied. The repreparation and computed tomographic (CT) scanning of its holotypic skull (MACN-Pv 18185) allowed a detailed description of its anatomy. Among the novel anatomical data presented here, we provide the first tridimensional complete reconstruction of a rhynchosaurid palate. This structure is extremely complex in comparison to the plesiomorphic archosauromorph condition, and more data on other rhynchosaurid palates are needed to shed light on the sequence of character states acquisition that resulted in this peculiar anatomy. The presence of a pair of septomaxillae in H. sanjuanensis is the first report of these bones in a rhynchosaurid, and this new information allowed to identify septomaxillae also in the hyperodapedontines Teyumbaita sulcognathus and Hyperodapedon mariensis. The better understanding of the skull anatomy of the holotype of H. sanjuanensis is a first step towards an improved knowledge of the morphology and taxonomy of the South American rhynchosaurs.