A taxonomic revision and cranial description of Terrestrisuchus gracilis (Archosauria, Crocodylomorpha) from the Upper Triassic of Pant-y-Ffynnon Quarry (southern Wales)

Non-crocodyliform crocodylomorphs, formerly referred to the informal group ‘Sphenosuchia’, are the earliest known crocodylomorph precursors of extant crocodylians. They are therefore crucial for our understanding of early crocodylian evolution and the origin of typical crocodylian characteristics, such as the formation of a secondary palate, complex cranial pneumaticity, and a reinforced braincase. Terrestrisuchus gracilis, known from the Upper Triassic fissure fill deposits of Pant-y-Ffynnon in southern Wales, is represented by almost 200 specimens, comprising articulated, partially associated, and isolated remains. In this contribution we provide a taxonomic revision of Terrestrisuchus gracilis and redescribe its cranial anatomy in detail, based in part on novel micro-computed tomography data. The posterior skull region is extensively pneumatized as indicated, among other features, by large pre- and postcarotid recesses on the parabasisphenoid, and a large pneumatic cavity in the articular of the mandible. In contrast, the quadrate forms only a small, unfused contact with the prootic, suggesting that complex pneumatization of the postorbital region pre-dated the co-ossification of the quadrate and braincase in Crocodylomorpha. Terrestrisuchus gracilis preserves an ossified basihyal, which represents the first occurrence of this bone in non-avemetatarsalian archosaurs. Finally, we show that Terrestrisuchus gracilis was probably cathemeral (i.e. active in a range of light levels), based on a phylogenetic flexible discriminant analysis of the relative dimensions of the sclerotic ring and orbit.