Abstract
We present a new phylogeny, based on DNA sequences of mitochondrial and nuclear genes, for frogs of the genus Osteocephalus with emphasis in the Osteocephalus buckleyi species complex. Genetic, morphologic, and advertisement call data are combined to define species boundaries and describe new species. The phylogeny shows strong support for: (1) a basal position of Osteocephalus taurinus + Osteocephalus oophagus, (2) a clade containing phytotelmata breeding species, and (3) a clade that corresponds to the Osteocephalus buckleyi species complex. Our results document a large proportion of hidden diversity within a set of populations that were previously treated as a single, widely distributed species, Osteocephalus buckleyi. Individuals assignable to Osteocephalus buckleyi formed a paraphyletic group relative to Osteocephalus verruciger and Osteocephalus cabrerai and contained four species, one of which is Osteocephalus buckleyi sensu stricto and three are new. Two of the new species are shared between Ecuador and Peru (Osteocephalus vilmae sp. n. and Osteocephalus cannatellai sp. n.) and one is distributed in the Amazon region of southern Peru (Osteocephalus germani sp. n.) We discuss the difficulties of using morphological characters to define species boundaries and propose a hypothesis to explain them.
Highlights
The Upper Amazon region has the highest alpha diversity of amphibians in the World with several sites exceeding 100 species in less than 10 km2 (Bass et al 2010)
Recent reviews (Jungfer 2010; 2011; Moravec et al 2009; Ron et al 2010) imply that the O. buckleyi species complex consists of nine species: O. buckleyi, O. cabrerai, O. carri, O. duellmani Jungfer 2011, O. festae, O. inframaculatus (Boulenger 1882), O. mutabor Jungfer and Hödl 2002, O. verruciger and an undescribed species sister to O. verruciger
Because most of our sampling was restricted to Ecuador and Peru, it is likely that there are even more species than found in our study. These results highlight the need to carry out large-scale genetic surveys of Amazonian amphibians to achieve a more realistic understanding of their diversity and evolution
Summary
The Upper Amazon region has the highest alpha diversity of amphibians in the World with several sites exceeding 100 species in less than 10 km (Bass et al 2010) These figures may vastly underestimate the total diversity as shown by the discovery of large numbers of cryptic species with the use of genetic markers (e.g., Fouquet et al 2007; Funk et al 2011; Padial and De la Riva 2009; Ron et al 2006). Its first large scale review was carried out by Trueb and Duellman (1971) who examined the morphology of specimens from seven countries and concluded that the O. buckleyi complex (excluding O. verruciger Werner 1901) consisted of a single, morphologically variable and widely distributed species They synonymized O. cabrerai, O. carri (Cochran and Goin 1970), and O. festae (Peracca 1904) under O. buckleyi. A phylogeny based on mitochondrial DNA revealed strong support for the O. buckleyi complex as well as paraphyly in O. verruciger and O. buckleyi (Ron et al 2010)
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