Taxonomy and phylogeny of the Aetosauria (Archosauria: Pseudosuchia) including a new species from the Upper Triassic of Arizona

Abstract

Aetosaurians are a clade of pseudosuchian archosaurs that were globally dispersed during the Late Triassic Epoch. Aetosaurians are characterized by a suite of osteoderms that covered much of the body. These osteoderms are commonly recovered as fossils and possess characteristic surface ornamentation that can be diagnostic for taxa. The abundance of these osteoderms and the ease of identification have made aetosaurians ideal index taxa for Late Triassic biostratigraphy. Of special interest are specimens from South and North America and Europe that have been assigned to the genus Stagonolepis, which have been utilized for correlation of continental sedimentary units and to approximately date the timing of important biotic events. New finds have called the synonymy of these Stagonolepis-like specimens into question, jeopardizing their ability to serve as biochronological markers. Detailed examination of all of the specimens assigned to Stagonolepis robertsoni demonstrates that all of these specimens do not represent the same species. The South American material is assigned to the genera Aetosauroides, Aetobarbakinoides, and Polesinesuchus; the European material to Stagonolepis; and the North American material to Calyptosuchus, Adamanasuchus, and a newly recognized taxon, Scutarx deltatylus. Scutarx deltatylus can be differentiated from other aetosaurians by the presence of a strongly raised, triangular boss, on the posteromedial corner of the paramedian osteoderms. Scutarx deltatylus also preserves the first good skull material from a Stagonolepis-like aetosaur from North America. A dorsoventrally thickened skull roof and an anteroposteriorly short parabasisphenoid further demonstrate the distinctness of this material from that of South America and Europe. A detailed phylogenetic analysis of all known aetosaurians further demonstrates the distinctness of these taxa. This new expanded analysis of 28 taxa and 83 characters recovers Aetosauroides scagliai as the sister taxon to all other aetosaurians. Stagonolepis robertsoni from Scotland does not clade with Stagonolepis olenkae from Poland. Calyptosuchus wellesi is the sister taxon to a clade consisting of Scutarx deltatylus and Adamanasuchus eisenhardtae. However, distribution of autapomorphies across these taxa precludes them from being synonymized. As a result the Stagonolepis-like aetosaurs cannot be used for global scale correlations of Upper Triassic strata, but do appear to be of utility for regional correlations, in particular those between the Chinle Formation and Dockum Group in the American Southwest.