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Phylogenetic Relationships of the South American Doradoidea (Ostariophysi: Siluriformes)

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Phylogenetic Relationships of the South American Doradoidea (Ostariophysi: Siluriformes) Neotropical Ichthyology, 12(3): 451-564, 2014 Copyright © 2014 Sociedade Brasileira de Ictiologia DOI: 10.1590/1982-0224-20120027 Phylogenetic relationships of the South American Doradoidea (Ostariophysi: Siluriformes) José L. O. Birindelli A phylogenetic analysis based on 311 morphological characters is presented for most species of the Doradidae, all genera of the Auchenipteridae, and representatives of 16 other catfish families. The hypothesis that was derived from the six most parsimonious trees support the monophyly of the South American Doradoidea (Doradidae plus Auchenipteridae), as well as the monophyly of the clade Doradoidea plus the African Mochokidae. In addition, the clade with Sisoroidea plus Aspredinidae was considered sister to Doradoidea plus Mochokidae. Within the Auchenipteridae, the results support the monophyly of the Centromochlinae and Auchenipterinae. The latter is composed of Tocantinsia, and four monophyletic units, two small with Asterophysus and Liosomadoras, and Pseudotatia and Pseudauchenipterus, respectively, and two large ones with the remaining genera. Within the Doradidae, parsimony analysis recovered Wertheimeria as sister to Kalyptodoras, composing a clade sister to all remaining doradids, which include Franciscodoras and two monophyletic groups: Astrodoradinae (plus Acanthodoras and Agamyxis) and Doradinae (new arrangement). Wertheimerinae, new subfamily, is described for Kalyptodoras and Wertheimeria. Doradinae is corroborated as monophyletic and composed of four groups, one including Centrochir and Platydoras, the other with the large-size species of doradids (except Oxydoras), another with Orinocodoras, Rhinodoras, and Rhynchodoras, and another with Oxydoras plus all the fimbriate-barbel doradids. Based on the results, the species of Opsodoras are included in Hemidoras; and Tenellus, new genus, is described to include Nemadoras trimaculatus, N. leporhinus and Nemadoras ternetzi. Due to conflicting hypotheses of the phylogenetic position of Acanthodoras, Agamyxis, and Franciscodoras, these are considered as incertae sedis in Doradidae. All suprageneric taxa of the Doradoidea are diagnosed based on synapomorphic morphological characteristics. Uma análise filogenética com base em 311 caracteres morfológicos é apresentada para a maioria das espécies de Doradidae, todos os gêneros de Auchenipteridae e representantes de outras 16 famílias de Siluriformes. A hipótese de consenso entre as seis árvores mais parcimoniosas corrobora o monofiletismo do grupo Sul-americano Doradoidea (Doradidae mais Auchenipteridae), bem como o de Doradoidea mais a família Africana Mochokidae. O clado de Sisoroidea mais Aspredinidae é considerado como grupo-irmão de Doradoidea mais Mochokidae. Dentre os Auchenipteridae, os resultados corroboraram o monofiletismo de Centromochlinae e Auchenipterinae. Esta última inclui Tocantinsia e outros quarto grupos monofiléticos, um com Asterophysus e Liosomadoras, um com Pseudotatia e Pseudauchenipterus, e outros dois maiores com os demais gêneros da subfamília. Dentre os Doradidae, a análise de parcimônia recuperou Wertheimeria como grupo-irmão de Kalyptodoras, formando um clado irmão de todos os demais doradídeos, que incluem Franciscodoras e dois grupos: Astrodoradinae (mais Acanthodoras e Agamyxis) e Doradinae (arranjo novo). Wertheimerinae, subfamília nova, é descrita para os gêneros Kalyptodoras e Wertheimeria. Doradinae é corroborada como monofilética e composta por quatro grupos de espécies, o primeiro com Centrochir e Platydoras, o segundo com os doradídeos de grande porte (exceto Oxydoras), o terceiro com Orinocodoras, Rhinodoras e Rhynchodoras, e o quarto com Oxydoras e os doradídeos de barbilhões fimbriados. Com base nos resultados, Opsodoras é considerado como sinônimo júnior de Hemidoras; e Tenellus, gênero novo, é descrito para Nemadoras trimaculatus, N. leporhinus e Nemadoras ternetzi. Devido à existência de hipóteses conflitantes para a posição filogenética de Acanthodoras, Agamyxis e Franciscodoras, estes são considerados como incertae sedis em Doradidae. Todos os grupos supra-genéricos de Doradoidea são diagnosticados com base em caracteres morfológicos sinapomórficos. Key words: Systematics, Taxonomy, Neotropical, Catfishes. Departamento de Biologia Animal e Vegetal, Universidade Estadual de Londrina. Caixa Postal 10.001, 86057-970 Londrina, PR, Brazil. [email protected]. 451 452 Phylogeny of Doradoidea Introduction discovered and the classification of the species of both families changed many times. Nevertheless, during the last Siluriformes comprises the most morphologically diverse, three decades, our understanding about the taxonomy and and geographically widespread order of Ostariophysi, with phylogenetic relationships of the species of Doradoidea has approximately 36 families, 477 genera, and 3088 valid species greatly improved. A review of the literature concerning the (Ferraris, 2007). Although most species are restricted to systematics of Doradoidea, and of each of its families, is freshwaters, some are tolerant of brackish waters, and one presented below. family occurs almost exclusively in marine environments The goals of the present study are to provide evidence (Nelson, 2006). for the phylogenetic position of the Doradoidea within Within the Siluriformes, the superfamily Doradoidea Siluriformes, to propose a hypothesis of phylogenetic comprises two Neotropical families: Doradidae and relationships among genera of the Auchenipteridae and Auchenipteridae. The species of Doradoidea range in length Doradidae, and to propose a new classification for Doradoidea. from a few centimeters (e.g., Physopyxis, Gelanoglanis) to a little more than one meter (e.g., Oxydoras). Most species are Systematic history of the Doradoidea. In the tenth edition of exclusive to freshwaters and occur in the largest river basins the Systema Naturae, Linnaeus (1758: 304, 307) described in South America, such as: Amazonas, La Plata, São Francisco, two genera to accommodate 15 species of catfish: Silurus, Parnaíba, Mearim, Orinoco, Magdalena, Maracaibo, with S. asotus, S. glanis, S. aspredo, S. mystus, S. anguillaris, Essequibo, and coastal rivers of Guyana, French Guiana, and S. batrachus, S. undecimalis, S. militaris, S. catus, S. clarias, Suriname. They are distinguished from other catfishes by S. ascita, S. costatus (=Platydoras costatus), S. callichthys, having a wide well-developed cephalic shield that is S. cataphractus (=Acanthodoras cataphractus), and the continuous to the dorsal-fin spine, posterior cleithral process monotypic Loricaria, with L. cataphracta. In the same usually large, and an infranuchal ligament (= ligament between volume, Linnaeus (1758: 238) described another catfish, posterior nuchal plate and rib of sixth vertebra; Birindelli et Acipenser plecostomus (=Hypostomus plecostomus, al., 2009; Birindelli et al., 2012). Loricariidae), among species of sturgeons (Acipenseridae). The species of Doradidae are popularly known as thorny Eight years later, in the 12th edition of the Systema Naturae, or talking catfishes (international community); cuiú-cuiús, Linnaeus (1766: 508) described Silurus inermis (=Ageneiosus bacus, botoados, armados, or rebecas (Brazil); and sierra or inermis), S. felis, S. galeatus (=Trachelyopterus galeatus), bagre hueso (Argentina, Bolivia, Colombia, Ecuador, S. cous, S. carinatus (=Doras carinatus), S. fasciatus, S. Paraguay, Peru, Uruguay, Venezuela). They are readily bagre, and reallocated A. plecostomus to Loricaria. Although distinguished among catfishes by having a single row of modern zoological nomenclature dates back to Linnaeus, his midlateral scutes, each usually with a single posteriorly classification was based on Peter Artedi’s (1738) oriented thorn. Vono & Birindelli (2007) described aspects of Ichthyologiae (Jordan, 1917). From Linnaeus to the end of the natural history of the monotypic Wertheimeria, and the 18th century, only a few studies described species and summarized published information on the ecology of other genera of Siluriformes. The most meaningful additions to thorny catfishes. Quagio-Grassiotto et al. (2011) provided Doradoidea were: Cataphractus Edwards (for former Silurus detailed descriptions and figures of sperm morphology in ten callichthys, S. cataphractus, and S. costatus), and Silurus species of doradids. nodosus Bloch (=Pseudauchenipterus nodosus). The species of Auchenipteridae are popularly known as In the 19th century, Bloch & Schneider (1801) proposed a driftwood, zamora or slope-head catfishes (international classification that grouped all species of fishes that were community); carataí, fidalgo, mandubé, palmito, judeu, then known into five classes. In one of the three orders of ximbé, cangati, or cachorro-de-padre (Brazil); and manduví, Octopterygii, they placed the species of Cataphractus and zamora, docella, or novia (Argentina, Bolivia, Colombia, Loricaria among non-catfishes; and in one of the three orders Ecuador, Paraguay, Peru, Uruguay, Venezuela). They are readily of Heptapterygii, they placed Silurus and Platystacus. distinguished among catfishes by exhibiting great sexual Lacepède (1803) described several genera of catfishes, dimorphism, in which mature males have the anal fin modified including Doras and Ageneiosus. Lacepède (1803) assigned for insemination. The ecology and biology of most Silurus nodosus and S. galeatus in Pimelodus, S. costatus, auchenipterids are not well documented. Large-sized species and S. carinatus in Doras; Ageneiosus armatus (junior of
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