Despite recent efforts to accelerate exploration and species description, the diversity of high Andean frogs remains highly underestimated. We report high levels of species diversity in direct-developing frogs or terraranas inhabiting the wet puna and adjacent cloud forests of the Amazonian versant of the Andes in Bolivia and Peru. Descriptive evidence of external morphology, distribution patterns and molecular phylogenetic analyses support the existence of nine unnamed species in two clades, which represents a 30% increase in species diversity for those clades. The relationships of these species and their relatives in Holoadeninae are tested using nuclear and mitochondrial genes for 159 terminals representing the 11 genera in this subfamily and 25 species of previously unknown relationships. Our results corroborate species monophyly in all but three cases and support the monophyly of al...

This study was partially funded by projects CLG2008-04164 and CLG2011-30393 of the Spanish government (PI: I. De la Riva), and the International Cooperation Project funded by the Consejo Nacional de Ciencia, Tecnologia e Innovacion Tecnologica (CONCYTEC, Peru) throught Fondo Nacional de Desarrollo Cientifico, Tecnologico y de Innovacion Tecnologica (FONDECYT, Peru) and the Ministerio de Ciencia, Tecnologia e Innovacion Productiva (MINCyT, Argentina) (PI. Juan C. Chaparro).

We infer species relationships within Lynchius, a frog genus with four species distributed along the paramos and cloud forests of the Andes of northern Peru and southern Ecuador, and assess species diversity in light of comparative analyses of anatomical traits and inferred relationships. Phylogenetic analyses rely on ~7000 base pairs of mtDNA and nuDNA sequences aligned using similarity-alignment and treealignment and optimized under maximum likelihood and parsimony criteria. Inferred relationships place Lynchius as the sister group of the widespread genus Oreobates and this clade as the sister group of the high Andean genus Phrynopus. Our analyses corroborate the dissimilar species Lynchius simmonsi as part of this clade and place it as the sister group of the remaining species of Lynchius. Parsimony and maximum likelihood analyses differ in the internal relationships of Lynchius with ...