THE EARLY EVOLUTION OF ARCHOSAURS: RELATIONSHIPS AND THE ORIGIN OF MAJOR CLADES

Archosaurs have a nearly 250 million year record that originated shortly after the Permian-Triassic extinction event and is continued today by two extant clades, the crocodylians and the avians. The two extant lineages exemplify two bauplan extremes among a diverse and complex evolutionary history, but little is known about the common ancestor of these lineages. Renewed interest in early archosaurs has led to nearly a doubling of the known taxa in the last 20 years. This study presents a thorough phylogenetic analysis of 80 species-level taxa ranging from the latest Permian to the early part of the Jurassic using a dataset of 412 characters. Each terminal taxon is explicitly described and all specimens used in the analysis are clearly stated. Additionally, each character is discussed in detail and nearly all of the character states are illustrated in either a drawing or highlighted on a specimen photograph. A combination of novel characters and comprehensive character sampling has bridged previously published analyses that focus on particular archosauriform subclades. A well-resolved, robustly supported consensus tree (MPTs = 360) found a monophyletic Archosauria consisting of two major branches, the crocodylian-line and avian-line lineages. The monophyly of clades such as Ornithosuchidae, Phytosauria, Aetosauria, Crocodylomorpha, and Dinosauria is supported in this analysis. However, phytosaurs are recovered as the closest sister taxon to Archosauria, rather than basal crocodylian-line archosaurs, for the first time. Among taxa classically termed as "rauisuchians," a monophyletic poposauroid clade was found as the sister taxon to a group of paraphyletic "rauisuchians" and monophyletic crocodylomorphs. Hence, crocodylomorphs are well nested within a clade of "rauisuchians," and are not more closely related to aetosaurs than to taxa such as Postosuchus. Basal crocodylomorphs such as Hesperosuchus and similar forms ("Sphenosuchia") were found as a paraphyletic grade leading to the clade Crocodyliformes. Among avian-line archosaurs, Dinosauria is well supported. A monophyletic clade containing Silesaurus and similar forms is well supported as the sister taxon to Dinosauria. Pterosaurs are robustly supported at the base of the avian line. A time-calibrated phylogeny of Archosauriformes indicates that the origin and initial diversification of Archosauria occurred during the Early Triassic following the Permian-Triassic extinction. Furthermore, all major basal archosaur lineages except Crocodylomorpha were established by the end of the Anisian. Early archosaur evolution is characterized by high rates of homoplasy, long ghost lineages, and high rates of character evolution. These data imply that much of the early history of Archosauria has not been recovered from the fossil record. Not only were archosaurs diverse by the Middle Triassic, but they had nearly a cosmopolitan biogeographic distribution by the end of the Anisian.