South American electric knifefishes of the genus Archolaemus (Ostariophysi, Gymnotiformes): undetected diversity in a clade of rheophiles

Neotropical electric knifefishes of the family Sternopygidae previously considered to represent a single relatively geographically widespread and morphologically variable species, Archolaemus blax, were analysed and found to represent a complex of five species, four of which are new to science. A fifth undescribed species from the Rio Sao Francisco basin outside the previous known range of the genus was identified. Recognized species of Archolaemus are: A. blax, previously thought to occur in the Rio Araguari, Rio Branco, Rio Tapajos, Rio Tocantins, and Rio Xingu, but which instead proved to be endemic to the Rio Tocantins; Archolaemus ferreirai sp. nov. from the Rio Mucajai and Rio Uraricoera in the north-eastern portions of the Amazon basin; Archolaemus janeae sp. nov. of the Rio Xingu and the upper Rio Tapajos, both southern tributaries of the mainstream Amazon; Archolaemus luciae sp. nov. of the Rio Trombetas, Rio Jari, and Rio Tapajos basins of the eastern Amazon, and the independent Rio Araguari draining into the Atlantic Ocean north of the mouth of the Amazon; Archolaemus orientalis sp. nov. of the Sao Francisco basin in eastern Brazil; and Archolaemus santosi sp. nov. of the Rio Jamari in the south-western portion of the Amazon basin. The phylogenetic placements of Archolaemus and the recently described genus Japigny relative to the other members of the Eigenmanninae are discussed. A series of synapomorphies for Archolaemus are proposed and a hypothesis of the relationships within that genus is advanced. Rheophily of all members of Archolaemus is discussed, with the genus found to be the most specious clade within the Gymnotiformes living primarily in high-energy settings. The reported anterior projection of the dentary teeth in A. blax was found to be a consequence of postmortem displacement.