The Pristimantis trachyblepharis species group, a clade of miniaturized frogs: description of four new species and insights into the evolution of body size in the genus

Species richness in the genus Pristimantis is underestimated due to the existence of morphologically cryptic species. This is worsened by the low sampling effort and the lack of studies using genetic markers. Here, we use molecular and morphological data to determine the phylogenetic relationships of a clade of Pristimantis distributed throughout montane tropical forests in the eastern Andes, from central Ecuador to northern Perú. We name this clade the Pristimantis trachyblepharis species group. Our results show that it comprises nine species, of which four are formally described and five are new. Four of these undescribed species are formally described here. The group is composed of miniaturized species, such as Pristimantis nanus sp. nov., currently the smallest known species of the genus and the smallest vertebrate in Ecuador. As a first approach to understanding the evolutionary origin and implications of body-size reduction in Pristimantis, we here study the phylogenetic signal and evolutionary trends of body size within the genus. We also provide the first record of P. aquilonaris in Ecuador and we show, for the first time, the phylogenetic position of P. albujai, P. aquilonaris, P. minimus and P. trachyblepharis, which are also members of the P. trachyblepharis species group.