Sistemática filogenética dos sapos-cuidadores aromobates e mannophryne (dendrobatoidea : aromobatidae : aromobatinae) inferida mediante evidência total

Abstract

Together, Aromobates and Mannophryne (Aromobatinae) include 39 recognized species distributed along the north of the Cordillera de los Andes and Cordillera de la Costa in Colombia, Venezuela, and Trinidad and Tobago. Previous studies supported the monophyly of these genera, their sister relationship, and elucidated the phylogenetic position of some of their species; however, these studies were based in an incomplete taxon sampling. Thus, the phylogenetic relationships of 41 % of the species in Aromobatinae remain unknown and the species diversity within both genera is not properly assessed. Herein we inferred the evolutionary relationships of Aromobatinae including for the first time all their recognized species. The inference was based on a parsimony total evidence analysis of DNA sequences of 31 mitochondrial and nuclear genes (analyzed under tree-alignment) and 383 phenotypic characters (188 of which are novel) sampled for 400 aromobatin and outgroup terminals. The ingroup is represented by a remarkably expanded taxonomic and geographic sampling, which includes 260 terminals representing the 39 recognized species and several putative new species from near 100 localities covering almost the entire geographic ranges of Aromobates and Mannophryne. Nine Aromobates species, as well as “Colostethus” caribe, “Prostherapis” dunni, and “Phyllobates” mandelorum, were included in the analyses only based on phenotypic data. The analyses rendered a well-resolved phylogeny, with Aromobates and Mannophryne as sister and reciprocally monophyletic. Novel and highly unexpected phylogenetic positions were discovered for “C.” caribe, “P.” dunni, and “P.” mandelorum within Aromobatidae. A fourth species group is recognized within Mannophryne and the M. collaris group is redefined. The 12 species of Aromobates included in this study for the first time are corroborated as part of this genus. Numerous novel intrageneric relationships and at least seven putative new species were also discovered. Aromobates ericksonae is considered a junior synonym of A. mayorgai. A new species of Mannophryne is described based on morphological and bioacoustic evidence, M. herminae is redefined, and the importance of bioacoustics evidence in species delimitations is discussed. Finally, we identified for the first time, numerous unambiguously optimized phenotypic synapomorphies for all supraspecific taxa and species groups. In the course of the field work of our study, three long-time missing species were rediscovered, among them, Mannophryne neblina, which had not been seen since its discovery 70 years ago. Our results highlight the importance of increasing the taxon sampling in phylogenetic analyses to improve the understanding of the systematics and species diversity. Also demonstrates the relevance of total evidence phylogenetic inference, for example by including species only represented by phenotypic data. According to the results, an updated taxonomy is proposed for Aromobatidae, concordant with our optimal topologies.