Combined phylogenetic analysis of Pleurodema (Anura: Leptodactylidae: Leiuperinae).

The genus Pleurodema comprises 15 species distributed through the Neotropical region, from sea level up to 5000 m.a.s.l. A total-evidence analysis of Pleurodema is provided based on the parsimony criterion. The combined dataset included morphometric, phenotypic, and DNA evidence (34 taxa, 4441 characters). The parsimony analysis yielded one most-parsimonious tree. Pleurodema was recovered as a well-supported clade composed of two major subclades. One subclade has an identical topology to that of previous analyses, the P. brachyops Clade (P. alium, P. borellii, P. brachyops, P. cinereum, P. diplolister, and P. tucumanum). The other subclade includes the remaining nine species of the genus, exhibiting a topology different from that of previous studies. According to the present phylogeny, this second lineage is formed by the P. nebulosum Clade (P. guayapae + P. nebulosum), P. marmoratum, the re-defined P. thaul Clade (P. bufoninum, P. somuncurense, P. thaul) and the P. bibroni Clade (P. bibroni, P. cordobae, P. kriegi). The reproductive modes of Pleurodema represent a unique combination of features within Leiuperinae, including three egg-clutch structures, two types of amplexus, and lack of vocalization. Also, some species of Pleurodema have been considered fossorial, because they are capable of digging with their hind-limbs and remaining in self-made burrows during dry seasons. The evolution of characters associated with reproductive biology and fossoriality is discussed in light of the obtained results.