Taxonomic revision ofEoalligator(Crocodylia, Brevirostres) and the paleogeographic origins of the Chinese alligatoroids
Background.The primarily Neotropical distribution of living alligatoroids raises questions as to when and how the ancestors ofAlligator sinensismigrated to China. As phylogeny provides a necessary framework for historical biogeographic issues, determining the phylogenetic positions of the Chinese alligatoroids is a crucial step towards understanding global alligatoroid paleobiogeography. Besides the unnamed alligatoroids from the Eocene of Guangdong Province, three Chinese fossil taxa have been referred to Alligatoroidea:Alligator luicus,Eoalligator chunyiiandEoalligator huiningensis. However, none of these fossil taxa has been included in a phylogenetic analysis. The genusEoalligatorwas established to accommodateE. chunyiifrom Guangdong Province.E. huiningensisfrom Anhui Province was later erected as a second species, despite no distinctive similarities withE. chunyii. By contrast, the putative crocodylineAsiatosuchus nanlingensiswas established based on material from Guangdong Province, close to theE. chunyiispecimens geographically and stratigraphically. Furthermore, specimens ofA. nanlingensisandE. chunyiishare four distinctive characters, but display no evident differences. As a result, the taxonomic relationships of these three species require restudy.Methods.In this paper, all specimens ofE. chunyiiandE. huiningensisare reassessed in detail, and compared to specimens ofA. nanlingensis. Detailed re-descriptions and revised diagnoses are provided, and a cladistic analysis is carried out to assess the phylogenetic positions ofE. chunyii,E. huiningensisandA. nanlingensis.Results.The analysis recoversE. chunyiiandA. nanlingensisas sister taxa among basal Crocodylidae, whileP. huiningensisis posited as an alligatoroid. Two key characters support the monophylyE. chunyii+A. nanlingensis: sulcus within surangular, and anteroposteriorly oriented surangular-articular suture. The former character is unique toE. chunyiiandA. nanlingensisamong crocodyloids, although a smaller and more posteriorly positioned surangular fossa is known inDiplocynodon. Detailed comparisons show the two species to be synonymous, withE. chunyiias the junior synonym ofA. nanlingensisbased on page priority. BecauseE. chunyiiwas erected as the type species ofEoalligator, the genus is now invalid. We establish the new genusProtoalligatorto accommodate “Eoalligator”huiningensis, an alligatoroid whose exact phylogenetic position is uncertain. In particular,P. huiningensisretains primitive characters such as a lacrimal that extends further anteriorly than the prefrontal, and a notch at the premaxilla-maxilla suture. However,P. huiningensisalso appears to share one important derived character, a complete nasal bar, with alligators. Our taxonomic revisions imply that four alligatoroids are currently known from China, and these species must have dispersed from North America to Asia in more than one event.