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Rodrigo Antunes Caires Rodrigo Antunes Caires REVISÃO TAXONÔMICA, ANATOMIA ESQUELÉTICA E FILOGENIA DO GÊNERO Microphilypnus Myers, 1927 (TELEOSTEI: GOBIIFORMES: ELEOTRIDAE). Departamento de Zoologia Instituto de Biociências da Universidade de São Paulo São Paulo Abril/2012 Rodrigo Antunes Caires REVISÃO TAXONÔMICA, ANATOMIA ESQUELÉTICA E FILOGENIA DO GÊNERO Microphilypnus Myers, 1927 (TELEOSTEI: GOBIIFORMES: ELEOTRIDAE). Tese apresentada no Departamento de Zoologia do Instituto de Biociências da Universidade de São Paulo, para obtenção de título de Doutor em Ciências, na área de Zoologia. Orientador: José Lima de Figueiredo Departamento de Zoologia Instituto de Biociências da Universidade de São Paulo São Paulo Abril/2012 ii Caires, Rodrigo Antunes Revisão taxonômica, anatomia esquelética e filogenia do gênero Microphilypnus (Teleostei: Gobiiformes: Eleotridae) xiii + 265 pp., 57 fig. Tese (Doutorado) – Instituto de Biociências da Universidade de São Paulo. Departamento de Zoologia. 1. Taxonomia, 2. Teleostei. 3. Filogenia I. Universidade de São Paulo. Instituto de Biociências, Departamento de Zoologia Banca examinadora ______________________ _____________________ Prof(a).Dr(a). Prof(a). Dr(a). ______________________ _____________________ Prof(a).Dr(a). Prof(a). Dr(a). _____________________________ Prof. Dr. José Lima de Figueiredo Orientador iii AVISO Esta dissertação é parte dos requerimentos necessários à obtenção do título de doutor, área de Zoologia e, como tal, não deve ser vista como uma publicação no senso do Código Internacional de Nomenclatura Zoológica (apesar de disponível publicamente sem restrições). Dessa forma, quaisquer informações inéditas, opiniões e hipóteses, bem como nomes novos, não estão disponíveis na literatura zoológica. Pessoas interessadas devem estar cientes de que referências públicas ao conteúdo deste estudo, na sua presente forma, devem ser somente feitas com anuência prévia do autor. NOTICE This dissertation is a partial requirement for the Ph.D. degree in Zoology and, as such, should not be considered as a publication in the sense of the International Code of Zoological Nomeclatural (although it is available without restrictions). Therefore, any new information, opinions, and hypothesis, as well as new names, are not available in the zoological literature. Interested people are advised that any public reference to this study, in its current form, should only be done after previous acceptance of the author. iv Este trabalho é dedicado àqueles que, ao longo da minha vida, me amaram do jeito que sou e me tornaram uma pessoa melhor: Érika, Fátima, Fernanda, Ferro, José Gonzaga, Renato, Vanessa. v “No fim, você pode não salvar sua alma e sua vida pelo pensamento. Mas, se você pensa, o menor dos prêmios de consolação é o mundo”. Saul Bellow, A Aventuras de Augie March. “Linguagem é coragem: a habilidade de conceber um pensamento, de proferi-lo e assim torná-lo verdadeiro Salman Rushdie, Os Versos Satânicos. vi RESUMO No presente projeto, foram realizados estudos sobre a taxonomia, a anatomia esquelética e as relações filogenéticas de Microphilypnus Myers, 1927 (Teleostei: Gobiidae: Eleotrinae). Este gênero de peixes é muito comum em tributários da Bacia Amazônica e do Orenoco, mas até o momento não era bem conhecido em termos taxonômicos. Com base na revisão da literatura e o exame de vários exemplares, duas das três espécies nominais foram validadas, uma espécie nova e duas espécies não descritas foram descobertas: Microphilypnus ternetzi Myers (bacias dos rios Negro, Orenoco, Amazonas, Madeira, Tapajós e Tocantins) (M. amazonicus Myers é sinônimo júnior desta espécie), Microphilypnus macrostoma Myers (bacias do Negro e Orenoco) Microphilypnus acangaquara Caires & Figueiredo (Baixo Tapajós), Microphilypnus sp. 1 (Baixo Tapajós) e Microphilypnus sp. 2 (rio Aripuanã). O estudo da anatomia esquelética baseou-se em 17 exemplares diafanizados e corados; foi constatado que as estruturas ósseas são semelhantes às dos demais membros de Gobioidei, e é postulado que o processo de miniaturização nesta espécie ocorreu por neotenia. Para a análise filogenética, foram incluídas, além de todas as espécies conhecidas de Microphilypnus, táxons de Gobioidei das instituições CAS, INPA, MZUSP e USNM, totalizando 56 terminais e 145 caracteres morfológicos. A análise filogenética resultou em 43 árvores mais parcimoniosas, com 1072 passos, IC: 0,22 e IR: 0,57. As relações de Microphilypnus resgatadas foram (M. macrostoma (M. ternetzi (Microphilypnus sp. 1(M. acangaquara + Microphilypnus sp. 2))), indicando a possibilidade de especiação parapátrica neste gênero. No consenso estrito, Microphilypnus figurou em uma politomia com os demais gêneros de Eleotrinae, ainda que no consenso semi-estrito tenha surgido como mais proximamente relacionado a dois gêneros de água doce: Leptophilypnus, da América Central, e Philypnodon, da Austrália, sugerindo que invasão do gênero objeto de estudo em água doce pode ter sido antiga. Na hipótese filogenética de consenso estrito deste projeto, Eleotrinae figurou como um grupo não monofilético, com Gobiomorus constituindo um ramo mais basal; Gobiidae surgiu como grupo monifilético, mas as subfamílias Gobiinae, Gobionellinae e Gobiosomatini surgiram como agrupamentos polifiléticos. São discutidos caracteres diagnósticos e as relações dos gêneros de Eleotrinae estudados, bem como das subfamílias de Gobiidae, e estudos filogenéticos adicionais com base em dados morfológicos são propostos. vii ABSTRACT This project entails a study on taxonomy, skeletal anatomy and phylogeny of Microphilypnus Myers, 1927 (Teleostei: Gobiidae: Eleotrinae). Members from this genus are very common in tributaries of the Orinoco and the Amazon Basin, but remained poorly known in taxonomic terms. Two of the three nominal species have been validated, a new species and two undescribed species were discovered: Microphilypnus ternetzi Myers (Negro, Orinoco, Amazon, Madeira, Tapajos, and Tocantins basins) (M. amazonicus Myers is a junior synonym); Microphilypnus macrostoma Myers (Negro and Orinoco basins); Microphilypnus acangaquara Caires & Figueiredo (lower Tapajós); Microphilypnus sp. 1 (lower Tapajós) and Microphilypnus sp. 2 (Aripuanã River). The study of skeletal anatomy was based on 17 cleared and stained specimens; it was found that the bone structures are similar to those of other members of Gobioidei, and it is postulated that species of this genus miniaturized by neoteny. For phylogenetic analysis, all Microphilypnus species and Gobiodei taxa from the institutions CAS, INPA, MZUSP and USNM were examined, totaling 56 terminals and 145 morphological characters. Phylogenetic analysis resulted in 43 most parsimonious trees with 1072 steps, CI = 0.22 and RI: 0.57. The relationships among Microphilypnus species that were retrieved in cladogram were: (Microphilypnus macrostoma (M. ternetzi (Microphilypnus sp. 1 (M. acangaquara + Microphilypnus sp. 2))), indicating a possibly parapatric speciation in this genus. In strict consensus, Microphilypnus figured in a polytomy other eleotrins, but in the majority-rule consensus it was nested with two freshwater genera: Leptophilypnus, from the Central America, and Philypnodon, from Australia, suggesting that invasion of freshwater in this genus may have been ancient. In the phylogenetic hypothesis of this project, Eleotrinae figured as a non-monophyletic group, with Gobiomorus constituting a more basal branch; Gobiidae emerged as a monophyletic group, but not the subfamilies Gobiinae, Gobionellinae and Gobiosomatini. Diagnostic characters and the relationships of Eleotrinae and subfamilies of Gobiidae are discussed herein. Additional phylogenetic studies based on morphological data are also proposed. viii AGRADECIMENTOS Este projeto só pôde ser realizado graças ao auxílio de inúmeras pessoas, e eu gostaria de agradecer a todas elas pessoalmente, porém faltam espaço e habilidade discursiva da minha parte para retribuir à altura. Desde já, peço perdão por eventuais esquecimentos de qualquer ajuda que eu tenha recebido. Da mesma forma, você, leitor que estiver com essa tese em mãos, pode se sentir ao menos parcialmente responsável por sua realização, pois foi com seus impostos que eu recebi financiamento para desenvolver o projeto. Sou grato aos professores e curadores da Seção de Peixes do Museu de Zoologia da Universidade de São Paulo (MZUSP) por toda a ajuda. Em particular, agradeço a meu orientador, Prof. José Lima de Figueiredo (Zé Lima). Enquanto eu estive trabalhando com ele, o Zé Lima foi extremamente atencioso, prestativo, referência como professor e pesquisador, e um dos melhores amigos que eu tive o prazer de conhecer. Ele provavelmente não imagina o quanto contribuiu, de forma decisiva, para o aprimoramento deste projeto, e na minha vida acadêmica e pessoal. Aos Profs. Heraldo Britski, Naércio Menezes e Mário de Pinna, pelas valiosas conversas, favores e sugestões para que eu pudesse tentar construir esse projeto e escrevê-lo. Também sou agradecido a todos os ictiólogos que trabalham ou trabalharam no MZUSP enquanto eu estive lá, pela paciência, pela amizade, e por compartilharem comigo parte do grande conhecimento de que dispõem. São eles: Alberto Akama, Andrea Paixão, André Casas, André Netto-Ferreira, Ângela Zanata, Alexandre Marceniuk, Aléssio Datovo, Bárbara Calegari, Caio, Carine Chamon, Carolina Paiva, Cléber, Cristiano Moreira, Diego Vaz, Douglas Silva, Eduardo Baena, Fábio di Dario, Fernando D’Agosta, Fernando Jerep, Flávia Petean, Flávio Lima, George Mattox, Gizelani Guazelli,
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