Phylogeny of Geophagine cichlids from South America (Perciformes: Labroidei)

Abstract

Three new species of cichlid fishes of the genus Geophagus, part of the Neotropical subfamily Geophaginae, are described from the Orinoco and Casiquiare drainages in Venezuela. Phylogenetic relationships among 16 genera and 30 species of Geophaginae are investigated using 136 morphological characters combined with DNA sequences coding for the mitochondrial gene NADH dehydrogenase subunit 4 (ND4) and the nuclear Recombination Activating Gene 2 (RAG2). Data from previous studies are integrated with the new dataset by incorporating published DNA sequences from the mitochondrial genes cytochrome b and 16S and the microsatellite flanking regions Tmo-M27 and Tmo-4C4. Total-evidence analysis revealed that Geophaginae is monophyletic and includes eighteen genera grouped into two major clades. In the first clade, the tribe Acarichthyini (genera Acarichthys and Guianacara) is sister-group to a clade in which Gymnogeophagus, 'Geophagus' steindachneri, and Geophagus sensu stricto are sister to 'Geophagus' brasiliensis and Mikrogeophagus; all these are in turn sister-group to Biotodoma, Dicrossus and Crenicara. In the second clade, Satanoperca, Apistogramma (including Apistogrammoides), and Taeniacara are sister to Crenicichla and Biotoecus. Monophyly and significantly short branches at the base of the phylogeny indicate that genera within Geophaginae differentiated rapidly within a relatively short period. High morphological, ecological, and behavioral diversity within the subfamily suggest that geophagine divergence may be the result of adaptive radiation.