Phylogenetic analysis of Ceratophryidae (Anura: Hyloidea) including extant and extinct species

The Neotropical frog family Ceratophryidae is composed of wide-mouthed frogs with stout bodies. Living species of the family are consistently recovered as a monophyletic group, but with disparities among analyses regarding internal relationships. Ceratophryidae presents one of the richest fossil records in Anura. Nevertheless, phylogenetic analyses including both extant and extinct species are still scarce, and the position of fossils is persistently debated. In this sense, the systematics of the family has changed considerably in the last decade with the exclusion of Baurubatrachus pricei (Late Cretaceous), Beelzebufo ampinga (Late Cretaceous) and Wawelia gerholdi (early Miocene). Herein, a morphologically based phylogeny for Ceratophryidae, including living species (11 spp.) and fossil specimens (10 spp.), is used as a background to discuss the evolutionary history of the family and its classification. We phylogenetically placed Baurubatrachus pricei, Beelzebufo ampinga and Wawelia gerholdi as non-ceratophryids. We recovered a monophyletic Ceratophryidae: Lepidobatrachus and Ceratophrys form a clade, with Chacophrys as its sister group. Our analysis corroborates the C. cornuta and C. aurita groups. Among fossils, L. australis and C. sagani were recovered as valid species based on autapomorphies, and C. rusconii was found to be the sister of all Ceratophrys. Ceratophrys ensenadensis, C. ameghinorum, C. aurita NHMUK PV OR18895/6 and C. sagani belong to the C. aurita group. We also discuss homoplasies in Ceratophryidae, divergence-time estimates, and the evolution of ploidy and a dorsal shield in the family.