The family Characidae is a group of freshwater bony fishes that exhibits high species-level diversity and whose members inhabit parts of Texas, Mexico, and Central and South America. Thus far, morphological data have been of limited use in discerning relationships among subfamilies and incertae sedis genera of the family Characidae. In this study, DNA sequence data from GenBank were combined with new sequences for analyses under Bayesian and parsimony schemes. Sequences fell into four gene partitions, with three genes in the mitochondrial subset (12S, 16S, COI genes) and one gene in the nuclear subset (RAG2 gene). Inferred Bayesian and parsimony-based phylogenies reject the monophyly of certain groups (e.g., Astyanax, Hyphessobrycon, and Bryconamericus), do not reject the monophyly of others (e.g., Cheirodontinae and "clade A" of previous authors), and present new sister-group hypotheses (e.g., Brittanichthys sister to Paracheirodon). Sister to clade A is a lineage referred to herein as "clade B" which includes Exodon and exemplars from Cheirodontinae (the most basal lineage within clade B), Aphyocharacinae, Tetragonopterinae, and Characinae (excluding Gnathocharax). "Clade C" is sister to A+B and contains representatives of large incertae sedis genera (e.g., Hyphessobrycon, Hemigrammus), as well as members of Stethaprioninae. Unless certain other subfamilial names are to be disregarded, the use of Tetragonopterinae should continue to be restricted to species of Tetragonopterus because other genera previously referred to this subfamily grouped in clades A or C, quite distant from Tetragonopterus.
The systematics of the Glandulocaudinae is reviewed in detail and justification for the recognition of the group as a subfamily is discussed. The subfamily Glandulocaudinae consists of three genera: Lophiobrycon with one species plesiomorphic in some anatomical features but some others exclusively derived relative to the species in the other genera; Glandulocauda with two species intermediate in phylogenetic derivation; and Mimagoniates with seven species (one new), all more phylogenetically derived concerning their pheromone producing caudal-fin organs and with other anatomical characters presumably more derived than in the species of the other genera. Glandulocauda melanogenys Eigenmann, 1911, is considered a junior synonym of Hyphessobrycon melanopleurus Ellis, 1911. A replacement name, Glandulocauda caerulea Menezes & Weitzman, is proposed for G. melanopleura Eigenmann, 1911. Gland cells found in the caudal-fin organs of all species are histologically indistinguishable from club cells and probably secrete a pheromone during courtship. The club cells are associated with somewhat modified to highly derived caudal scales forming a pheromone pumping organ in the more derived genera and species. This subfamily is distributed in freshwaters of eastern and southern Brazil, Paraguay, and northeastern Uruguay. A sistemática de Glandulocaudinae é revista e a justificativa para o reconhecimento do grupo como subfamília discutida. A subfamília Glandulocaudinae consiste de três gêneros: Lophiobrycon, com uma espécie plesiomórfica com relação a alguns caracteres anatômicos, mas outros derivados e exclusivos em relação às espécies dos outros dois gêneros; Glandulocauda, com duas espécies intermediárias quanto à condição dos caracteres derivados; e Mimagoniates, com sete espécies (uma nova), todas filogeneticamente mais avançadas quanto às características dos órgãos da nadadeira caudal produtores de feromônio e outras características anatômicas presumivelmente mais derivadas do que nas espécies dos outros gêneros. Glandulocauda melanogenys Eigenmann, 1911, é considerado sinônimo junior de Hyphessobrycon melanopleurus Ellis, 1911. O nome Glandulocauda caerulea Menezes & Weitzman, é proposto em substiutição para G. melanopleura Eigenmann, 1911. Células glandulares encontradas nos órgãos da caudal são histologicamente indistinguíveis de "células club" e provavelmente secretam algum tipo de feromônio durante a corte. As "células club" são associadas a escamas da caudal pouco ou inteiramente modificadas e fazendo parte dos órgãos bombeadores de feromônio nas espécies e gêneros mais derivados. Esta subfamília distribui-se em ambientes de água doce do leste e sul do Brasil, no Paraguai e nordeste do Uruguai
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