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Pontifícia Universidade Católica Do Rio Grande Do Sul PONTIFÍCIA UNIVERSIDADE CATÓLICA DO RIO GRANDE DO SUL FACULDADE DE BIOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM ZOOLOGIA Taxonomy and Phylogeny of the Neotropical fish genus Hemibrycon Günther, 1864 (Ostariophysi: Characiformes: Characidae) Vinicius de Araújo Bertaco Orientador: Dr. Luiz Roberto Malabarba TESE DE DOUTORADO PORTO ALEGRE - RS - BRASIL 2008 SUMÁRIO Dedicatória.................................................................................................................................iii Agradecimentos.........................................................................................................................iv Resumo......................................................................................................................................vi Abstract.....................................................................................................................................vii Apresentação............................................................................................................................viii Capítulo I – Taxonomy and Phylogeny of the Neotropical fish genus of Hemibrycon Günther, 1864 (Ostariophysi: Characiformes: Characidae) ……………………………………………..1 Capítulo II – A new species of Hemibrycon (Teleostei: Characiformes: Characidae) from the río Ucayali drainage, Sierra del Divisor, Peru (publicado em 2007, Neotropical Ichthyology, 5(3): 251-257) ………………………………………………………………………..……..288 Conclusões Gerais………………………………………………………………………...…296 ii A Verônica, e a meus pais. iii Agradecimentos Ao meu Orientador, Prof. Dr. Luiz Roberto Malabarba, pela dedicação e excelente orientação, amizade e apoio durante todo o desenvolvimento deste trabalho. A minha Família, por me incentivar, apoiar, ajudar e mostrar o caminho da sabedoria e do sucesso e, principalmente, por me dar condições de chegar até aqui. Um agradecimento especial para os meus pais e minha avó, Wanda M. de Araújo (in memorian), para mim, exemplos de vida. Julinana Bertaco revisou parte do texto redigido em inglês. A Verônica C. Baumbach, pelo companheirismo, amor, apoio e incentivo durante todas as etapas deste estudo, e muita paciência... Aos pesquisadores do Laboratório de Ictiologia do Museu de Ciências e Tecnologia da PUCRS, Carlos A. S. de Lucena, Zilda Margarete S. Lucena e Roberto E. dos Reis, pelo grande apoio e dedicação em algumas discussões levantadas sobre taxonomia e filogenia de Characiformes nas aulas do curso de doutorado, e durante o desenvolvimento deste trabalho. Aos colegas do Laboratório de Ictiologia do Museu de Ciências e Tecnologia da PUCRS, Alexandre Cardoso, Cristian Cramer, Cristina Bührnheim, Edson Pereira, Fernando Jerep, Juliana Wingert, Ignacio Moreno, José Pezzi da Silva, Monica Rodrigues, Pablo Lehmann e Tiago Carvalho, pela colaboração em alguma etapa deste estudo. Ao Dr. Stanley Weitzman (USNM) pelas discussões e sugestões via e-mail, e pelo auxílio obtido para a visita de curta duração a coleção de peixes do National Museum of Natural History, Smithsonian Institution, Whashington D.C. Aos curadores ou responsáveis de coleções científicas pelo empréstimo do material para o estudo: Scott Schaefer e Barbara Brown (AMNH), Jonh Lundberg, Mark Sabaj-Perez e Kyle Luckenbill (ANSP), J. Fong, David Catania, e T. Iwamoto (CAS), Kevin Swagel, Marie Anny Rogers e Philip Willink (FMNH), Ivan Mojica (ICNMHN), Armando Ortega-Lara (IMNC), M. Retzer (INHS), A. Bently (KU), Donald Taphorn (MCNG), Sonia Fisch-Muller e A. Merguin (MHNG), Pablo Lehmann e Santiago Ayerbe (MHNUC), Guy Duhamel e Patrice Pruvost (MNHN), Hernan Ortega e Max Hidalgo (MUSM), Osvaldo Oyakawa, Flávio Lima e Pedro Holanda (MZUSP), Helmut Wellendorf e Ernst Mikschi (NMW), Sven Kullander e E. Ahlander (NRM), R. Winterbotton e E. Holm (ROM), Stanley Weitzman e Sandra Raredon (USNM), Isaäc Isbrücker (ZMA), e Jorgen Nielsen (ZMUC). A Andréa Thomaz (UFRGS) pelos cortes histológicos das gônadas de alguns espécimes de Hemibrycon, e ao Marco A. Azevedo (FZB) pelas imagens do MEV das gônadas e arcos branquiais de algumas espécies de Hemibrycon. iv Ao Carlos Santanna (INPA), Javier Maldonado (IAvHP) e Pedro Holanda (MZUSP) pela total assistência durante as minhas visitas aos museus de história natural Norte Americanos, Colombianos e Suíço, respectivamente. Ao Kevin Swagel (FMNH), Roberto Reis e Tiago Carvalho (MCP), Paulo Petry (MCZ) e a Monica Rodrigues pelo exame e fotos do material tipo de algumas espécies de Hemibrycon e Bryconamericus. A Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) pela concessão da bolsa de doutorado. A Pontifícia Universidade Católica do Rio Grande do Sul e Museu de Ciências e Tecnologia pela infra-estrutura e recursos oferecidos, sem os quais não seria possível a realização deste trabalho. Ao Programa de Pós-Graduação em Zoologia desta universidade, pela oportunidade de realizar o curso. Ao Centro de Microscopia e Microanálises (CEMM) da PUCRS, pelas imagens de microscopia eletrônica de varredura dos dentes, arcos branquiais e gônadas de alguns espécimes de Hemibrycon. As instituições científicas estrangeiras ANSP, CAS, FMNH, MHNG e USNM pelo auxílio financeiro concedido para visitar as respectivas coleções de peixes. A todos que me ajudaram direta ou indiretamente para a concretização deste estudo, o meu MUITO OBRIGADO! v Resumo As espécies do gênero Hemibrycon Günther (1864) são revisadas e redescritas com base na análise do material tipo e exemplares adicionais. Hemibrycon é considerado monofilético e relacionado ao Clado A sensu Malabarba & Weitzman (2003) com base na análise de 123 caracteres e 45 táxons. No total, 21 espécies de Hemibrycon são reconhecidas, incluindo cinco novas espécies. Hemibrycon n. sp. 1 da bacia superior do río Ucayali, Peru; Hemibrycon n. sp. 2 de pequenos rios de drenagens costeiras do Mar do Caribe, Colômbia; Hemibrycon n. sp. 3 e Hemibrycon n. sp. 4 da porção média da bacia do río Magdalena, Colômbia, e Hemibrycon n. sp. 5 da bacia superior do río Madre de Dios, Peru. Estas espécies distinguem-se, principalmente, pelo padrão de colorido e número de raios ramificados da nadadeira anal, de escamas da linha lateral e de dentes no maxilar. A área de distribuição do gênero é ampliada para a bacia do baixo rio Tocantins, Brasil, com a ocorrência de H. surinamensis nessa bacia, sendo o primeiro registro da espécie para este país. A posição taxonômica de Bryconamericus decurrens e Hemibrycon orcesi é discutida. Novos sinônimos são propostos para algumas espécies de Hemibrycon: H. coxeyi é sinônimo júnior de H. polyodon, H. carrilloi é sinônimo júnior de H. dariensis e H. guppyi é sinônimo júnior de H. taeniurus. É fornecida uma chave de identificação para as espécies examinadas de Hemibrycon. vi Abstract The species of the genus Hemibrycon Günther (1864) are reviewed and redescribed based on their type series and additional specimens. Hemibrycon is hypothesized to be monophyletic and related to Clade A genera sensu Malabarba & Weitzman (2003) based on the analysis of 123 characters including 45 taxa. A total of 21 species are recognized in Hemibrycon, including five new species. Hemibrycon n. sp. 1 from the upper río Ucayali drainage, Peru; Hemibrycon n. sp. 2 from small rivers of the Caribbean Sea coastal drainages of Colombia; Hemibrycon n. sp. 3 and Hemibrycon n. sp. 4 from the middle río Magdalena basin, Colombia, and Hemibrycon n. sp. 5 from the upper río Madre de Dios, Peru. These species are distinguished among themselves mainly in the color pattern and number of branched anal-fin rays, lateral line scales, and maxillary teeth. The known geographical distribution of the genus is enlarged reaching the lower rio Tocantins basin, Brazil, with the occurrence of H. surinamensis in this area, the first record of the species for that country. The taxonomic status of Bryconamericus decurrens and Hemibrycon orcesi is discussed. New synonymies are proposed for some species of Hemibrycon: H. coxeyi is a junior synonym of H. polyodon, H. carrilloi is a junior synonym of H. dariensis, and H. guppyi is a junior synonym of H. taeniurus. An identification key to the examined species of Hemibrycon is provided. vii Apresentação Esta tese é apresentada conforme as Normas do Programa de Pós-Graduação em Zoologia da Pontifícia Universidade Católica do Rio Grande do Sul - PUCRS. A tese está estruturada em forma de dois artigos científicos redigidos em inglês, um já publicado na Neotropical Ichthyology. É apresentado um resumo em português. As tabelas e figuras estão no final do texto do Capítulo I. O periódico escolhido para a publicação desse capítulo foi Neotropical Ichthyology, revista da Sociedade Brasileira de Ictiologia com periodicidade trimestral. Os artigos científicos foram redigidos conforme as instruções para os autores do periódico escolhido. Neste estudo foi realizada uma revisão e redescrição das espécies incluídas no gênero Hemibrycon Günther, 1864 com base na análise do material tipo e exemplares adicionais e uma análise filogenética das relações de Hemibrycon com representantes do Clado A proposto por Malabarba & Weitzman (2003) e outros gêneros de Characidae (Capítulo I). O gênero Hemibrycon é diagnosticado com base em sinapomorfias relacionadas a caracteres osteológicos e de morfologia externa e relacionado aos gêneros do Clado A sensu Malabarba & Weitzman (2003). Um total de 21 espécies é reconhecido para o gênero Hemibrycon, incluindo cinco novas espécies. Hemibrycon n. sp. 1 da bacia superior do rio Ucayali, Peru; Hemibrycon n. sp. 2 de pequenos rios de drenagens
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