Dealing with homoplasy: osteology and phylogenetic relationships of the bizarre neobatrachian frog Baurubatrachus pricei from the Upper Cretaceous of Brazil

The hyperossified frog Baurubatrachus pricei Báez & Peri 1989 from the Maastrichtian Serra da Galga Member of the Marília Formation is described in detail, as preparation of the type and only known specimen revealed significant features, particularly of the pectoral and pelvic girdles. This species is rediagnosed on the basis of the combination of plesiomorphic and derived character states, including two unique traits: cranial roof with round openings that might have contained the tympanic membrane completely circumscribed by ornamented dermal bone, and scapula bearing a conspicuous crest deflected ventrally to form a deep basin on its leading edge. Since its discovery it was suggested that Baurubatrachus might be a relative of the South American ceratophryids, a phylogenetic placement endorsed by recent analyses. In order to test this hypothesis considering all the available information, we conducted several maximum parsimony analyses under different weighting schemes and topological constraints, scoring 143 characters for 71 extant and extinct anuran taxa. Our taxonomic sampling included species with well-ossified dermatocrania as well as less ossified members of main neobatrachian clades to explore the impact of hyperossification, which frequently drives groupings based on homoplastic features. We also assessed the phylogenetic signal provided by cranial and postcranial partitions. Although we recovered a monophyletic Ceratophryidae repeatedly, Baurubatrachus was not related with this nobleobatrachian group but associated with the calyptocephalellid australobatrachians, although with weak support. Other possible phylogenetic placements are also discussed, as well as microhabitat and habits, taking into account both anatomical and geological data.