Reassessment of the enigmatic crocodyliform " Goniopholis" paulistanus Roxo, 1936: Historical approach, systematic, and description by new materials

Abstract

The Crocodyliformes are the most represented vertebrate clade in the Upper Cretaceous sequences of the Bauru Group, Paraná Basin. However, some of the species described have an uncertain taxonomic status and phylogenetic position. For instance, “Goniopholis” paulistanus has been assigned as a nomem dubium, due to its description being based on scarce material. The “ G”. paulistanus specimens (i.e. teeth and a left tibia) were discovered in two different localities in São Paulo state: Mirandópolis and Valparaíso municipalities; where the upper interval of the Adamantina Formation (Early Maastrichtian of Bauru Group) crops out. Revisiting these specimens, we observed multicrenulated teeth in middle dentary toot- row, a remarkable feature only shared with teleosaurids Machimosaurus hugii (Upper Jurassic of Laurasia) and M. rex (Lower Cretaceous of Tunisia). This apomorphy was also recognized in new material from the Alfredo Marcondes municipality (Presidente Prudente Formation), which are here also referred to “ G”. paulistanus. We recognized the teeth of “ G.” paulistanus as the lectotype, however the tibia cannot be assigned to a species as it was not collected in association with the teeth. We performed a phylogenetic analysis with a data matrix composed of 388 characters and 86 taxa, analyzed in TNT. The strict consensus tree recovered Neosuchia and Ziphosuchia (Notosuchia + Sebecia) within Mesoeucrocodylia. The species “G” paulistanus is valid, as a distinct and new genus within Sebecia, in a polytomy with Barreirosuchus, Pepesuchus, Itasuchus and Peirosaurus, forming the clade Itasuchidae. Stolokrosuchus is the sister taxon to Itasuchidae, the sister group of all other Sebecia (Peirosauridae (Mahajangasuchidae + Sebecidae and taxa affinis)). The clades Ziphosuchia, Sebecia and Itasuchidae are here redefined, and we find the last two clades to be more closely related to terrestrial notosuchids than to semiaquatic neosuchians.