Molecular insights into the phylogeny of Mormyriform fishes and the evolution of their electric organs

In this report we generate a partial phylogeny of the mormyriform fishes using mitochondrial DNA sequences from twelve species of mormyriforms belonging to five genera. Electric organs and electric organ discharges are also examined. We have sequenced and aligned 373 bases from the mitochondrial 12S rRNA and 559 bases from the 16s rRNA from fourteen species of the superorder Osteoglossomorpha. Two non-mormyriform genera were used as outgroups. Three phylogenetic methods generated concordant partial phylogenies for these fish. Our analysis focuses on the genus Brienomyrus, which is a heterogeneous clade with at least eleven nominal species. Six morphs from Gabon had distinctive EODs but were morphologically 'cryptic' in that they all had the brachyistius-like body morphology. DNA analysis fully supports the EOD data that the six morphs represent distinct clades. The group from Gabon is monophyletic, while B. brachyistius from West Africa is a separate lineage. B. niger, a second distinct lineage, is a sister group to the six species from Gabon. Petrocephalus is the sister group of all the genera of the subfamily Mormyrinae so far analyzed, thereby confirming previous osteological results. Gymnarchus niloticus is the sister group of the family Mormyridae, also confirming an earlier phylogenetic hypothesis based on morphology. The molecular data adds polarity to electric organ characteristics. Stalkless electrocytes appear to be primitive. Petrocephalus, with non-penetrating stalked electrocytes innervated on the posterior side, represents an ancestral state for the Mormyridae, while Marcusenius, Brienomyrus and Gnathonemus with penetrating-stalked electrocytes, represent the apomorphic condition. Two species with doubly-penetrating electrocytes innervated on the posterior side may represent a transitional stage. At least two species of Brienomyrus appear to have reverted to non-penetrating stalked electrocytes, possibly through paedomorphosis.