The two major lineages of extant archosaurs, crocodylians and avians, diverged in the Triassic, but the details and timing of this event are incompletely understood. Fragmentary and phylogenetically uninformative specimens, in addition to poor temporal constraints on rock units from the Early and Middle Triassic, typify obstacles in identifying early archosaurs. This paper re-describes the partial skeleton of the only known specimen of Xilousuchus sapingensis Wu, 1981 from the Early Triassic Heshanggou Formation in north-central China. Originally assigned to the non-archosaurian archosauriform clade Proterosuchidae, an extensive phylogenetic analysis posits X. sapingensis as a crown-group archosaur within Suchia, thus making this taxon the unequivocally oldest known member of Archosauria. The age and phylogenetic position of X. sapingensis indicate that many archosaurs, including all major clades of non-archosaurian archosauriforms, the avianline, ornithosuchids, aetosaurs and paracrocodylomorph lineages, must have diverged by the end of the Early Triassic. X. sapingensis is part of a possible clade of sail-backed poposauroids that were common components of archosaur assemblages during the Early to Middle Triassic.
Largocephalosaurus polycarpon Cheng et al. 2012a was erected after the study of the skull and some parts of a skeleton and considered to be an eosauropterygian. Here we describe a new species of the genus, Largocephalosaurus qianensis, based on three specimens. The new species provides many anatomical details which were described only briefly or not at all in the type species, and clearly indicates that Largocephalosaurus is a saurosphargid. It differs from the type species mainly in having three premaxillary teeth, a very short retroarticular process, a large pineal foramen, two sacral vertebrae, and elongated small granular osteoderms mixed with some large ones along the lateral most side of the body. With additional information from the new species, we revise the diagnosis and the phylogenetic relationships of Largocephalosaurus and clarify a set of diagnostic features for the Saurosphargidae Li et al. 2011. Largocephalosaurus is characterized primarily by an oval supratemporal fenestra, an elongate dorsal ‘rib-basket’, a narrow and elongate transverse process of the dorsal vertebrae, and the lack of a complete dorsal carapace of osteoderms. The Saurosphargidae is distinct mainly in having a retracted external naris, a jugal–squamosal contact, a large supratemporal extensively contacting the quadrate shaft, a leaf-like tooth crown with convex labial surface and concave lingual surface, a closed dorsal ‘rib-basket’, many dorsal osteoderms, a large boomerang-like or atypical T-shaped interclavicle. Current evidence suggests that the Saurosphargidae is the sister-group of the Sauropterygia and that Largocephalosaurus is the sister-group of the Saurosphargis–Sinosaurosphargis clade within the family.
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