Taxonomy, systematics and geographic distribution of ground frogs (Alsodidae, Eupsophus): a comprehensive synthesis of the last six decades of research.

Abstract

The genus Eupsophus (ground frogs) inhabits exclusively the temperate forests of southern South America (Chile and Argentina). The current delimitation of the genus was reached in the late 1970s, when only two species were recognized, but since then the number of described species steadily increased, reaching a maximum of 11 by 2012. Subsequent studies that applied explicit species delimitation approaches decreased the number of species to six in 2017 and raised it again to 11 the following year, including an undescribed putative species. Despite these taxonomic changes, the two species groups traditionally recognized, roseus and vertebralis, have been maintained. Another recent contribution to the taxonomy of the genus was the explicit recognition of the extremely high level of external phenotypic variation exhibited by species of the roseus group, which undermines the utility of some diagnostic characters. Here we provide a critical review of the extensive taxonomic and systematic literature on the genus over the last six decades, to examine the evidence behind the recurrent taxonomic changes and advances in its systematics. We also update and complete a 2017 review of geographic information, provide additional qualitative observations of external characters commonly used in the diagnoses of species of the roseus group, and reassess the phylogenetic position of a putative new species from Tolhuaca (Chile), which was not included in the last species delimitation study. The present review shows that: 1) there is no congruence between the patterns of phenotypic and genetic/phylogenetic differentiation among species of both groups; 2) in the roseus group, the intraspecific variation in some external characters is as high as the differences described among species; 3) there is little morphological and bioacoustic differentiation within species groups, and inconsistencies in the chromosomal evidence at the genus level; 4) under the latest taxonomic proposal (2018), species of the roseus group still lack consistent and reliable diagnoses and their distribution limits are poorly defined; and 5) the population from Tolhuaca represents an additional undescribed species under the most recent taxonomic framework. Finally, we discuss the implications of these findings for the taxonomy and biogeography of the genus, pointing out some areas that require further research to understand their patterns and processes of diversification.