THE BRAINCASE AND ENDOCAST OF PARRINGTONIA GRACILIS, A MIDDLE TRIASSIC SUCHIAN (ARCHOSAUR: PSEUDOSUCHIA)

The evolution of the braincase and brain of early pseudosuchians through to the earliest crocodylomorphs is poorly understood given the paucity of specimens, lack of well-preserved material, and lack of consensus on the phylogenetic relationships of the major clades of Pseudosuchia. Here, we describe three differently sized braincases diagnosable as belonging to the archosaur Parringtonia gracilis from the Middle Triassic Manda Beds of Tanzania. One of them, a nearly complete and exceptionally preserved braincase, possesses a complex set of character states typically present in early-diverging Pseudosuchia, but it also possesses nearly all of the character states previously hypothesized to represent synapomorphies exclusive to aetosaurs and crocodylomorphs. Details of the internal anatomy and a reconstructed endocast were obtained through high-resolution computed tomographic data and show a high degree of conservation of brain architecture across Pseudosuchia, in stark contrast to the extensive shifts observed within Theropoda to crown Aves. Integrating cranial and postcranial data into a phylogenetic analysis, we find Parringtonia gracilis to be closely related to a Revueltosaurus-Aetosauria clade at the base of Suchia and distantly related to crocodylomorphs. No matter which pseudosuchian clade is the most closely related to crocodylomorphs, we deduce that homoplasy in the braincase appears to be common across Pseudosuchia.