The fossil record and phylogeny of South American horned frogs (Anura, Ceratophryidae)

Abstract

South American horned frogs (Ceratophryidae), with their large heads, wide gapes and fang-like teeth, are among the most charismatic, best-known and well-studied neobatrachian anurans. The family comprises 12 extant species with hyperossified skulls and has a relatively rich fossil record, particularly in the Pampas, which dates back to the late Miocene. However, several records have been overlooked in recent summaries, and many taxonomic assignments remain indeterminate or are questionable and have yet to be tested within a quantitative phylogenetic framework. Here we provide a complete up-to-date survey of the palaeontological record of Ceratophryidae, including some remarkable new records. We also tested their systematic position through comprehensive phylogenetic analyses based on osteological data, providing several synapomorphies for all relevant nodes. Finally, we discuss these integrated data in relation to divergence time estimates, and propose a set of fossil calibrations that provide hard minimum bounds for crown-group Ceratophryidae and the subclades within it, and illuminate the acquisition of polyploidy within the group.