The diversity of Triassic South American sphenodontians: a new basal form, clevosaurs, and a revision of rhynchocephalian phylogeny

Rhynchocephalians are a group of lizard-like diapsid reptiles that were very diverse during the Mesozoic but are now restricted to a single extant genus in New Zealand. Recent cladistic analyses have revealed two major clades, Eusphenodontia and the more crownward Neosphenodontia, but relationships of individual taxa have remained difficult to determine because of missing data and an unrevised data matrix. Here we drastically revise the established data matrix on rhynchocephalians by reassessing, evaluating, and adding new characters and operational taxonomic units, differing from any previous analyses in our goal to consider all known rhynchocephalians. In addition, we describe a new genus and species of an early eusphenodontian taxon from the Norian of southern Brazil, with a unique mosaic of plesiomorphic and apomorphic traits, and we re-examine the craniodental anatomy of the eusphenodontian Clevosaurus brasiliensis with µCT imaging, revealing a unique form of acrodonty amongst rhynchocephalians.Fil: Chambi Trowell, Sofia A. V.. University of Bristol; Reino UnidoFil: Martinelli, Agustín Guillermo. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; ArgentinaFil: Whiteside, David I.. University of Bristol; Reino UnidoFil: Romo de Vivar Martínez, Paulo Rodrigo. Universidade Federal do Rio Grande do Sul; BrasilFil: Bento Soares, Marina. Universidade Federal do Rio Grande do Sul; BrasilFil: Schultz, Cesar. Universidade Federal do Rio Grande do Sul; BrasilFil: Gill, Pamela G.. University of Bristol; Reino UnidoFil: Benton, Michael J.. University of Bristol; Reino UnidoFil: Rayfield, Emily J.. University of Bristol; Reino Unid

A New Clevosaurid from the Triassic (Carnian) of Brazil and the Rise of Sphenodontians in Gondwana

The early evolution of lepidosaurs is marked by an extremely scarce fossil record during the Triassic. Importantly, most Triassic lepidosaur specimens are represented by disarticulated individuals from high energy accretion deposits in Laurasia, thus greatly hampering our understanding of the initial stages of lepidosaur evolution. Here, we describe the fragmentary remains of an associated skull and mandible of Clevosaurus hadroprodon sp. nov., a new taxon of sphenodontian lepidosaur from the Late Triassic (Carnian; 237–228 Mya) of Brazil. Referral to Sphenodontia is supported by the combined presence of a marginal dentition ankylosed to the apex of the dentary, maxilla, and premaxilla; the presence of ‘secondary bone’ at the bases of the marginal dentition; and a ventrally directed mental process at the symphysis of the dentary. Our phylogenetic analyses recover Clevosaurus hadroprodon as a clevosaurid, either in a polytomy with the Late Triassic to Early Jurassic Clevosaurus and Brachyrhinodon (under Bayesian inference), or nested among different species of Clevosaurus (under maximum parsimony). Clevosaurus hadroprodon represents the oldest known sphenodontian from Gondwana, and its clevosaurid relationships indicates that these sphenodontians achieved a widespread biogeographic distribution much earlier than previously thought.Fil: Hsiou, Annie S.. Universidade de Sao Paulo; BrasilFil: Nydam, Randall L.. Midwestern University; Estados UnidosFil: Simões, Tiago R.. University of Alberta; Canadá. Harvard University; Estados UnidosFil: Pretto, Flávio A.. Universidade Federal de Santa Maria; BrasilFil: Onary, Silvio. Universidade de Sao Paulo; BrasilFil: Martinelli, Agustín Guillermo. Universidade Federal do Rio Grande do Sul; Brasil. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; ArgentinaFil: Liparini, Alexandre. Universidade Federal de Sergipe; BrasilFil: Romo de Vivar Martínez, Paulo Rodrigo. Universidade Federal do Rio Grande do Sul; BrasilFil: Soares, Marina. Universidade Federal do Rio Grande do Sul; BrasilFil: Schultz, Cesar. Universidade Federal do Rio Grande do Sul; BrasilFil: Caldwell, Michael Wayne. University of Alberta; Canad

New rhynchocephalian specimen in the Late Triassic of southern Brazil and comments on the palatine bone of Brazilian rhynchocephalians

Triassic rhynchocephalians from South America are relatively sparse, based on one taxon from the Norian of Argentina (i.e., Sphenotitan leyesi) and three records from the Late Carnian (i.e., indet. taxon) and Norian (i.e., Clevosaurus brasiliensis and indet. taxon) of Brazil. We described here a new occurrence of rhynchocephalians from a new Late Carnian locality of southern Brazil, based on an isolated palatine with teeth, referred to that group due to the morphological similarity whit the palatines of the other taxa of the same group. The specimen comes from the municipality of Vale do Sol, collected within a layer with abundant material of Hyperodapedon sp. (Rhynchosauria), which is referred to the Hyperodapedon Assemblage Zone (Late Carnian) of the Candelária Sequence, Santa Maria Supersequence. The isolated palatine here described cannot be referred to any hitherto known species, but it provides an additional record for the few putative ones of Carnian age in South America. Moreover, we figure and describe the palatine anatomy of Clevosaurus brasiliensis, based on two specimens, which have relevant data poorly addressed in previous contributions.Fil: Romo de Vivar Martínez, Paulo Rodrigo. Universidade Federal do Rio Grande do Sul; BrasilFil: Martinelli, Agustín Guillermo. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; ArgentinaFil: Paes Neto, Voltaire D.. Universidade Federal do Rio Grande do Sul; BrasilFil: Scartezini, Camila A.. Universidade Federal do Rio Grande do Sul; BrasilFil: Lacerda, Marcel B.. Universidade Federal do Paraná; BrasilFil: Rodrigues, Carlos N.. Museu Municipal Aristides Carlos Rodrigues; BrasilFil: Soares, Marina Bento. Universidade Federal do Rio Grande do Sul; Brasi