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Characiformes: Characidae: Cheirodontinae) PONTIFÍCIA UNIVERSIDADE CATÓLICA DO RIO GRANDE DO SUL PROGRAMA DE PÓS-GRADUAÇÃO EM ZOOLOGIA REVISÃO TAXONÔMICA E FILOGENIA DA TRIBO COMPSURINI (CHARACIFORMES: CHARACIDAE: CHEIRODONTINAE) FERNANDO CAMARGO JEREP PORTO ALEGRE, 2011 PONTIFÍCIA UNIVERSIDADE CATÓLICA DO RIO GRANDE DO SUL FACULDADE DE BIOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM ZOOLOGIA REVISÃO TAXONÔMICA E FILOGENIA DA TRIBO COMPSURINI (CHARACIFORMES: CHARACIDAE: CHEIRODONTINAE) FERNANDO CAMARGO JEREP ORIENTADOR: DR. LUIZ ROBERTO MALABARBA TESE DE DOUTORADO PORTO ALEGRE - RS - BRASIL 2011 Aviso Este trabalho é parte integrante dos requerimentos necessários à obtenção do título de doutor em Zoologia, e como tal, não deve ser vista como uma publicação no senso do Código Internacional de Nomenclatura Zoológica (artigo 9) (apesar de disponível publicamente sem restrições) e, portanto, quaisquer atos nomenclaturais nela contidos tornam-se sem efeito para os princípios de prioridade e homonímia. Desta 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, somente devem ser feitas com aprovação prévia do autor. Notice This work is a partial requirement for the PhD degree in Zoology and, as such, should not be considered as a publication in the sense of the International Code of Zoological Nomenclature (article 9) (although it is available without restrictions) therefore, any nomenclatural acts herein proposed are considered void for the principles of priority and homonymy. Therefore, any new information, opinions, and hypotheses, 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. SUMÁRIO Dedicatória ............................................................................................................ ii Agradecimentos ..................................................................................................... iv Resumo .................................................................................................................. vii Abstract ................................................................................................................. viii Apresentação ......................................................................................................... ix Capítulo I - Systematics of Compsurini Malabarba, Weitzman & Burns (Characiformes: Characidae) .................................................................................. 1 Capítulo II - Revision of the genus Compsura Eigenmann, 1915 (Characidae: Cheirodontinae: Compsurini), with description of two new species from the Amazon basin, Brazil ............................................................................................. 229 Capítulo III - Revision of the genus Macropsobrycon Eigenmann, 1915 (Characidae: Cheirodontinae: Compsurini) ............................................................ 273 Capítulo IV - A revisionary study of the trans-Andean Compsurini (Characidae: Cheirodontinae): description of a new genus and review of the Saccoderma Schultz, 1944 ......................................................................................................... 306 Capítulo V - Description of three genera and six new species of Compsurini (Characidae: Cheirodontinae) from the upper Amazon basin .................................. 363 Capítulo VI - Redescription of Leptobrycon xinguensis (Géry, 1972), new combination, and phylogenetic relationships and redefinition of Leptobrycon (Characiformes: Characidae) .................................................................................. 419 Conclusões Gerais.................................................................................................. 452 i Dedicatória Dedico este trabalho a todos piabólogos e às piabas piabas difíceis de serem triadas piabas difíceis de serem contadas piabas dificeis de serem identificadas piabas mal-vistas a primeira vista, e escondidas no passado interespecificamente aglomeradas em lotes empoeirados abarrotadas como sardinhas, em álcool negligenciado sofrendo de descamação precoce e perda de raio por errôneo manuseio e formol mal graduado piaba minúscula, pequena, média e grande, coletadas em rio, riacho, arroio, lagoa e ponte piabas que migram da nascente ao estuário piabas que acabam em lojas de aquário com colorido prateado, dourado, vermelho, ou azulado que sustenta garça, ciclídeo, bagre, traíra e botuado piaba com pseudotímpano e sem mancha umeral com pedúnculo arqueado ou sem linha lateral piaba sem dente na maxila e com escamas na caudal duas fileiras no dentário e i+9 na dorsal piaba que não inova, piaba que não varia, piaba desprovida de autapomorfia, piabas que politomizam hipóteses e monotonizam resultados homoplasicamente agrupadas em clados mal suportados piaba de isca, piaba que belisca piaba que mama mas não fisga. ii P Q Rabindranath Tagore iii Agradecimentos Ao meu orientador Luiz R. Malabarba, por todos os ensinamentos e orientação nos momentos de devaneio e ignorância, pela disposição, entusiasmo, paciência e convivência agradável. Mas principalmente por expandir meu horizonte científico. Ao programa de Pós-graduação em Zoologia da PUCRS e seus alicerces, os coordenadores Roberto E. Reis e Júlio César Bicca-Marques, e as secretárias Luiza e Luana, que foram exemplos de eficiência e presteza em cada etapa deste curso de doutorado. Aos grandes professores ictiólogos Carlos A. Lucena, Zilda Margarete Lucena, Roberto E. Reis e Édson Pereira, cujo precioso conhecimento me foi transmitido por difusão facilitada no dia a dia de convivência no laboratório, nas conversas durante o cafezinho, jogos de bola, coletas e almoços (aê tinha pastel?). Ao curador Carlos A. Lucena por toda assessoria nos empréstimos de material, conserto de lupa, troca de lâmpadas, acesso à internet, e compartilhamento de erva de chimarrão. Aos amigos laboratoriais que por aqui estão ou por aqui passaram: Alexandre Cardoso, Alexandre Scharkasnky, Christian Cramer, Cristina Bührnheim, Cristina Oliveira, Bárbara Calegari, Professor Edson Pereira, Fernanda Meyer sobrinha da dona da Armelin, Fernando “Cumpadi” R. Carvalho, Héctor Paraguayo, Ignacio “Iggy” Moreno, Janine “Molusco” Arruda, Juliano Romanzini, Lautaro, Marco A. Azevedo, Mariangeles Arce, Rodrigo Quevedo, Tiaguinho Carvalho, Vinicius A. Bertaco, Vivi e financiador Carlins Jacobs, e Zé Pezzi. E aos mais novos, Maria Laura, Natália, Beatriz, Rachel, Jonas e Filipe. Às três “fias” ictiólogas, Bá, Mari e Vivis, pela ajuda e parcerismo ao longo destes quatro anos. Aos amigos e colegas laboratoriais da UFRGS: Ana Paula Dufech, Andrea Thomaz, profª Clarice Fialho, Circe, Clayton Fukakusa, Dudu Machado, Fernanda Weiss, Fernando R. de Carvalho, Giovana Lagemann, Giovanni Neves, Guilérme Formooosa Correa, Júlia Giora, Juliano Ferrer dos Santos, Juliana Wingert, Tatiana Dias, Vinicius Bertaco e Vinicius Lampert. Um agradecimento especial ao piabeiros Ademir, Paulo Nunes, Cumpadi de VB e VAB plantador de grama e melancia, pelos momentos memoráveis nas coletas e prosas sobre iv piaba; um agradecimento especial pela ajuda na expedição em busca dos Compsurini, do rio Guaporé ao rio Araguaia. Aos amigos e colegas ictiólogos de outras terras pelas conversas, úteis e inúteis, porém ambas agradáveis, e troca de experiências que tive durante estes quatro anos, durante visitas a coleções, coletas e disciplinas: Alexandre Ribeiro, André Casas, André Netto-Ferreira, Andréa Paixão, Clarianna Silva, Cláudio Zawadzki, Cristiano Moreira, Flávio Lima, Henrique Varella, Javier Maldonado, José Birindelli, Katiane Ferreira, Karinão, Leandro Sousa, Manuela Marinho, Marcelo Britto, Marcelo Rocha, Pedro Hollanda, Rafaela Ota, Ricardo Benine, Mateus Soares, Marcel Cavallaro de Lucélia, Aléssio Datovo de Divinolândia, Fernandinho Lagosta, Jupi do Tocantins, e Willian China Massaharu. Aos amigos e professores da Universidade Estadual de Londrina, pelo contínuo aprendizado que se estende aos dias de hoje: Dr. Oscar A. Shibatta, Dr. Mário Orsi, Dra. Lúcia Giuliano, e Dr. Sirlei Bennemann. E aos amigos londrinenses Édson, Cido, e Wanner. À professora Dr. Irani Quagio-Grassiotto (UNESP) pelas imagens de ultraestrutura dos espermatozóides de muitos Compsurini. Aos curadores e pesquisadores que ajudaram a realização deste trabalho, seja pelo empréstimo de material ou por sugestões pertinentes: Carla Pavanelli (NUPELIA), Carlos A. Lucena (MCP), Cláudio Oliveira (UNESP), Francisco Langeani (UNESP), Heraldo Britski (MZUSP), Jansen Zuanon (INPA), Lucia Py- Daniel (INPA), Marcelo Costa (UFRJ), Mônica Toledo-Piza Ragazzo (IBUSP), Naércio Menezes (MZUSP), Oscar A. Shibatta (UEL), Osvaldo T. Oyakawa (USP), e Paulo Buckup (MNRJ). For the loan of specimens and for the museum and technical support, I am thankful to Scott Schaefer (AMNH), Barbara Brown (AMNH), John Lundberg, Mark Sabaj Pérez (ANSP), David Catania, Jon D. Fong (CAS), Mary Anne Rogers (FMNH), Kevin Swagel (FMNH), Jose Ivan Mojica (ICNMHN), Lawrence M. Page (INHS), Marlis R. Douglas (INHS), Michael E. Retzer (INHS), Sonia Müller (MHNG), Richard Vari, Jerry Finan, Jeff Clayton (NMNH), R. Winterbottom (ROM), E. Holm (ROM), Larry Page (UF), and Doug Nelson (UMMZ). Special thanks to Richard P. Vari (NMNH) for supervising me during my
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