The systematic relationships and biogeographic history of ornithischian dinosaurs
The systematic relationships of taxa traditionally referred to as ‘basal ornithopods’ or ‘hypsilophodontids’ remain poorly resolved since it was discovered that these taxa are not a monophyletic group, but rather a paraphyletic set of neornithischian taxa. Thus, even as the known diversity of these taxa has dramatically increased over the past two decades, our knowledge of their placement relative to each other and the major ornithischian subclades remained incomplete. This study employs the largest phylogenetic dataset yet compiled to assess basal ornithischian relationships (255 characters for 65 species level terminal taxa). The resulting strict consensus tree is the most well-resolved, stratigraphically consistent hypothesis of basal ornithischian relationships yet hypothesized. The only non-iguanodontian ornithopod (=basal ornithopod) recovered in this analysis is Hypsilophodon foxii. The majority of former ‘hypsilophodontid’ taxa are recovered within a single clade (Parksosauridae) that is situated as the sister-taxon to Cerapoda. The Parksosauridae is divided between two subclades, the Orodrominae and the Thescelosaurinae. This study does not recover a clade consisting of the Asian taxa Changchunsaurus, Haya, and Jeholosaurus (=Jeholosauridae). Rather, the former two taxa are recovered as basal members of Thescelosaurinae, while the latter taxon is recovered in a clade with Yueosaurus near the base of Neornithischia.The endemic South American clade Elasmaria is recovered within the Thescelosaurinae as the sister taxon to Thescelosaurus. This study supports the origination of Dinosauria and the early diversification of Ornithischia within Gondwana. Neornithischia first arose in Africa by the Early Jurassic before dispersing to Asia before the late Middle Jurassic, where much of the diversification among non-cerapodan neornithischians occurred. Under the simplest scenario the Parksosauridae originated in North America, with at least two later dispersals to Asia and one to South America. However, when ghost lineages are considered, an alternate dispersal hypothesis has thescelosaurines dispersing from Asia into South America (via North America) during the Early Cretaceous, then back into North America in the latest Cretaceous. The latter hypothesis may explain the dominance of orodromine taxa prior to the Maastrichtian in North America and the sudden appearance and wide distribution of thescelosaurines in North America beginning in the early Maastrichtian. While the diversity of parksosaurids has greatly increased over the last fifteen years, a ghost lineage of over 40 myr is present between the base of Parksosauridae and Cerapoda, indicating that much of the early history and diversity of this clade is yet to be discovered. This new phylogenetic hypothesis provides a comprehensive framework for testing further hypotheses regarding evolutionary patterns and processes within Ornithischia.