Taxonomy of Probolodus Eigenmann, 1911 (Characiformes: Characidae) with description of two new species, and comments about the phylogenetic relationships and biogeography of the genus

Probolodus is a genus of tetras distributed along the coastal basins of southeastern Brazil, from Espirito Santo on the north, to Sao Paulo states. It was proposed by Eigenmann in 1911, to include the single species P. heterostomus. However, examining of material recently collected we identified two new species: P. oyakawai, new species, from the rio Ribeira de Iguape basin, and P. sazimai, new species, from the rio Itapemirim (Espirito Santo), and drainages of the rio Doce basin (Espirito Santo and Minas Gerais states). Therefore, the distribution of P. heterostomus was restricted to the rio Paraiba do Sul basin, southeastern Brazil. The genus Probolodus was redefined based on osteological characters, in particular details of oral dentition. Probolodus heterostomus differs from its congeners by a combination of meristic and osteological characters. An identification key for the species of the genus is presented. The possible evolutionary relationships of Probolodus with other characids, especially some lepidophagous taxa are discussed. Regarding the biogeography of the group, some geological events are hypothesized as responsible for the cladogenetic events among species of the genus.