Project description: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.

Project description:One of the most characteristic and abundant amphibian taxa of South American temperate forests is Eupsophus. The ten currently recognized species of the genus have been divided in two species groups, roseus and vertebralis, but most of them, eight, belong to the roseus group. Recent phylogeographic and phylogenetic studies have suggested that species diversity of the roseus group could be underestimated. An examination of the literature shows that species of the roseus group exhibit high levels of variation in their external characteristics, particularly those used as diagnostic characters, which compromises their taxonomy and hinders their field recognition. High levels of variation were also observed in several new populations of the roseus group discovered in southern Chile (36°-40°S), which could not be identified to the species level by their external characteristics. On the other hand, the literature reveals a scarse karyotype differentiation and a high bioacoustic uniformity among the species of the roseus group. We performed a Bayesian phylogenetic analysis using mitochondrial and nuclear genes to reevaluate the species diversity of the roseus group, including all the nominal species of Eupsophus and new populations. This analysis was complemented with three species delimitation approaches, General Mixed Yule Coalescent, multi-rate Poisson Tree Process and Automatic Barcode Gap Discovery. We favored a conservative delimitation of only four species for the roseus group, a result more consistent with the distribution of pairwise genetic distances, and the available chromosome and bioacoustic evidence. The four recognized lineages, which have nearly completely allopatric distributions, are named after the earliest nominal species that they include, but because high levels of phenotypic variation, they are not diagnosable by consistent differences in external morphology. We discuss the implications of this new proposal for the taxonomy and conservation of the genus, and the possible causes of the difficulty to estimate its species diversity.

Project description:The alsodid ground frogs of the Eupsophus genus are divided into two groups, the roseus (2n = 30) and vertebralis (2n = 28), which are distributed throughout the temperate Nothofagus forests of South America. Currently, the roseus group is composed by four species, while the vertebralis group consists of two. Phylogenetic relationships and species delimitation within each group are controversial. In fact, previous analyses considered that the roseus group was composed of between four to nine species. In this work, we evaluated phylogenetic relationships, diversification times, and species delimitation within the roseus group using a multi-locus dataset. For this purpose, mitochondrial (D-loop, Cyt b, and COI) and nuclear (POMC and CRYBA1) partial sequences from 164 individuals were amplified, representing all species. Maximum Likelihood (ML) and Bayesian approaches were used to reconstruct phylogenetic relationships. Species tree was estimated using BEAST and singular value decomposition scores for species quartets (SVDquartets). Species limits were evaluated with six coalescent approaches. Diversification times were estimated using mitochondrial and nuclear rates with LogNormal relaxed clock in BEAST. Nine well-supported monophyletic lineages were recovered in Bayesian, ML, and SVDquartets, including eight named species and a lineage composed by specimens from the Villarrica population (Bootstrap:>70, PP:> 0.99). Single-locus species delimitation analyses overestimated the species number in E. migueli, E. calcaratus, and E. roseus lineages, while multi-locus analyses recovered as species the nine lineages observed in phylogenetic analyses (Ctax = 0.69). It is hypothesized that Eupsophus diversification occurred during Mid-Pleistocene (0.42-0.14 Mya), with most species having originated after the Last Southern Patagonian Glaciation (0.18 Mya). Our results revitalize the hypothesis that the E. roseus group is composed of eight species and support the Villarrica lineage as a new putative species.

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Eupsophus altor

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