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Characiformes: Characidae) FERNANDA ELISA WEISS SISTEMÁTICA E TAXONOMIA DE HYPHESSOBRYCON LUETKENII (BOULENGER, 1887) (CHARACIFORMES: CHARACIDAE) Tese apresentada ao Programa de Pós-Graduação em Biologia Animal, Instituto de Biociências da Universidade Federal do Rio Grande do Sul, como requisito parcial à obtenção do Título de Doutora em Biologia Animal. Área de Concentração: Biologia Comparada Orientador: Prof. Dr. Luiz Roberto Malabarba Universidade Federal do Rio Grande do Sul Porto Alegre 2013 Sistemática e Taxonomia de Hyphessobrycon luetkenii (Boulenger, 1887) (Characiformes: Characidae) Fernanda Elisa Weiss Aprovada em ___________________________ ___________________________________ Dr. Edson H. L. Pereira ___________________________________ Dr. Fernando C. Jerep ___________________________________ Dra. Maria Claudia de S. L. Malabarba ___________________________________ Dr. Luiz Roberto Malabarba Orientador i Aos meus pais, Nelson Weiss e Marli Gottems; minha irmã, Camila Weiss e ao meu sobrinho amado, Leonardo Weiss Dutra. ii 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. iii Agradecimentos Ao meu orientador, Dr. Luiz R. Malabarba, pela orientação em todos os momentos dessa caminhada, pela disposição, entusiasmo e paciência. Ao Dr. Richard Vari, por toda assistência científica, pela cordialidade, pelo acolhimento recebido durante minha permanência em Washington, D.C., e pelo auxílio financeiro que possibilitou a extensão da minha permanência nos Estados Unidos. Ao Dr. John Lundberg (ANSP), pelo “grant” que propiciou minha visita na coleção ictiológica da Academy of Natural Sciences of Philadelphia. Ao Programa de Pós-graduação em Biologia Animal da UFRGS, e todo corpo docente pela infra estrutura e ensinamentos. Ao suporte financeiro do Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), que concedeu minha bolsa de doutorado, e bolsa de doutorado sandwich no exterior (procs. 140401/2009-1 e 201264/2011-0). Aos membros da comissão avaliadora, Dr. Edson H. L. Pereira, Dr. Fernando C. Jerep e Dra. Maria Claudia de S. L. Malabarba, pela disponibilidade em participar da análise desse trabalho. Aos colegas do laboratório de Ictiologia da UFRGS: Alice Hirschmann, Aline Vanin, Ana Paula Dufech, Andréa Maciente, Andrea Thomaz, Andréia Turcati, Andrei Langoni, Antonio Queiroz Lezama, Caroline Hartmann, Profª Dra. Clarice Fialho, Claudio Ricken, Clayton Kunio Fukakusa, Fernando R. de Carvalho, Guilherme Frainer, Ingrid Tossedo, João Paulo Santos, Julia Giora, Juliana Wingert, Juliano Ferrer dos Santos, Karina Tondato, Karine Orlandi Bonato, Laura Modesti Donin, Letícia Schönhofen Longoni, Luiz Artioli, Natália G. Berthier, Priscilla Caroline, Renato Bolson Dala Corte, Tatiana Schmidt Dias, Vinicius Bertaco e Vinicius Renner Lampert., pelo bom ambiente de trabalho e trocas de idéias. Um agradecimento especial a Antonio Queiroz Lezama por ajudar a resolver todas as dúvidas de filogenia; a Andréa Maciente e Juliana Wingert pela ajuda com as impressões da tese, enquanto eu estava longe; a Clayton Kunio Fukakusa e João Paulo Santos pela documentação fotográfica e artística, e pelos excelentes “Sábados Gourmet”, e finalmente à grande amiga Tatiana Schmidt Dias, pela diagramação, edicão de arte, apoio e amizade em todas as fases desse trabalho. iv Aos amigos e colegas ictiólogos de outras instituições: Adriana Saccol-Pereira, Andréa Paixão, Carlos Oliveira, Renata Bartolette, Tiago Carvalho e Vivianne Sant'Anna, pelas troca de experiências e visitas a coleções durante esses quatro anos. Aos curadores e pesquisadores que ajudaram na realização deste trabalho, seja através do empréstimo de material ou pelo suporte técnico em laboratório: Carlos Lucena (MCP), Claudia Machado (LAPAD), Francisco Langeani (UNESP), Luisa Sarmento-Soares (MM), Marco Aurélio Azevedo (FZB), Miriam Ghazzi, Sonia Buck (UFSC), Osvaldo T. Oyakawa (MZUSP) e Vinícius Abilhoa (MHCI), James Maclaine, Oliver Crimmen (BMNH), John Lundberg, Mark Sabaj Pérez (ANSP), David Catania, Jon D. Fong (CAS), Richard Vari, Jerry Finan, Jeff Clayton e Sandra Raredon (preparação de fotos e raio X) (NMNH), Karsten Hartel (MCZ) e Helmut Wellendorf (NMW). À Karen Lee, pela acolhida em Washington D.C. e Sasha Li, pela hospedagem em San Francisco (CA). Aos amigos que compartilhei bons momentos em terras americanas: Alessandro Bifi, Aline Staskowian Benetti, Dahiana Arcila, Danilo Pacheco Cordeiro, Lorian Cobra Straker e Rubens Coelho. Um super agradecimento especial ao “fiooooo” Fernando Jerep, por todo suporte técnico, científico e cultural, e a “companheira de lombas” Vivianne Sant'Anna. A minha grande amiga Rita, cuja presença constante foi responsável pela finalização dessa tese. Ao meu namorado, David Krancer, pela vivência na cultura americana, pelo companherismo, e por ter colaborado com o aprimoramento da minha fluência na Língua Inglesa “for free”. E por fim, pórem não menos importante, um agradecimento especial aos meus amados pais, Nelson e Marli, minha irmã Camila e meu sobrinho Leo, por toda força e incentivo constantes em todos os momentos, ao longo desse trajeto. v Sumário Lista de Tabelas........................................................................................................................ ix Lista de Figuras......................................................................................................................... xi Resumo................................................................................................................................... xiv Abstract.................................................................................................................................... xv Apresentação.............................................................................................................................. 1 Referências Bibliográficas......................................................................................................... 5 Capítulo 1 – Systematics of Hyphessobrycon luetkenii (Boulenger, 1887) (Characiformes: Characidae)................................................................................................................... 11 Abstract........................................................................................................................ 12 Resumo......................................................................................................................... 12 Key words.................................................................................................................... 13 Introduction.................................................................................................................. 13 Material and Methods................................................................................................... 16 Taxon sampling................................................................................................ 16 Institutional abbreviations................................................................................ 16 Cladistic methodology...................................................................................... 17 Results and Discussion................................................................................................. 18 Phylogenetic diagnoses for clades and terminal taxa....................................... 18 General discussion............................................................................................ 37 Comparative material....................................................................................... 39 Literature Cited............................................................................................................ 40 Appendix I. Data set of species analyzed in addiction to data set of Mirande (2009, 2010) and Carvalho (2011). Taxa are listed in alphabetic order. Polymorphisms are denoted as “z”, inapplicable data as “-” and, missing data as “?”................................ 69 vi Appendix II. List of characters including number of steps, and consistency and retention indices of each character in the final hypothesis herein proposed under implied weighting......................................................................................................... 72
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