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Revisão Taxonômica E Relações Interespecíficas De André Luiz Netto Ferreira REVISÃO TAXONÔMICA E RELAÇÕES INTERESPECÍFICAS DE LEBIASININAE (OSTARIOPHYSI: CHARACIFORMES: LEBIASINIDAE) Departamento de Zoologia Instituto de Biociências Universidade de São Paulo Outubro/2010 André Luiz Netto Ferreira REVISÃO TAXONÔMICA E RELAÇÕES INTERESPECÍFICAS DE LEBIASININAE (OSTARIOPHYSI: CHARACIFORMES: LEBIASINIDAE) Tese apresentada ao Instituto de Biociências da Universidade de São Paulo, para a obtenção de Título de Doutor em Ciências, na Área de Zoologia. Orientador: Mário C. C. de Pinna Departamento de Zoologia Instituto de Biociências Universidade de São Paulo Outubro/2010 Netto-Ferreira, André Luiz Revisão taxonômica e relações interespecíficas de Lebiasininae (Ostariophysi: Characiformes: Lebiasinidae) xix+460 pp., 107 fig. Tese (Doutorado) - Instituto de Biociências da Universidade de São Paulo. Departamento de Zoologia. 1. Sistemática 2. Neotropical 3. Teleostei 4. Cladística 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. Mário C. C. de Pinna Orientador 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). 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 dissertation 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 (although it is available without restrictions). 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. Àqueles que caminharam comigo, ainda que tenham se juntado tardiamente, nessa longa jornada. Em especial para minhas três pessoinhas queridas. v Their world is fashioned. Natural Science No wonder they don't understand. 1. Tide Pools 3. Permanent Waves When the ebbing tide retreats Science, like nature, Along the rocky shoreline, Must also be tamed It leaves a trail of tidal pools With a view towards its preservation. In a short-lived galaxy. Given the same Each microcosmic planet State of integrity, A complete society. It will surely serve us well. A simple kind mirror Art as expression, To reflect upon our own. Not as market campaigns All the busy little creatures Will still capture our imaginations. Chasing out their destinies. Given the same Living in their pools, State of integrity, They soon forget about the sea... It will surely help us along. Wheels within wheels in a spiral array, The most endangered species, A pattern so grand and complex, The honest man, Time after time we lose sight of the way, Will still survive annihilation. Our causes can't see their effects. Forming a world State of integrity, 2. Hyperspace Sensitive, open and strong. A quantum leap forward In time and in space, Wave after wave will flow with the tide, and The universe learned to expand. bury the world as it does. Tide after tide will flow and recede, leaving The mess and the magic, life to go on as it was... Triumphant and tragic, A mechanized world out of hand. Lee, Lifeson & Peart Computerized clinic For superior cynics Who dance to a synthetic band. In their own image, vi Resumo A subfamília Lebiasininae é tradicionalmente dividida em dois gêneros: Lebiasina e Piabucina. Historicamente, tal divisão vem sendo contestada devido à inconsistência dos caracteres que diferenciam os dois gêneros. Com o intuito de investigar as relações entre os Lebiasininae e a validade de Lebiasina e Piabucina, foram analisados 59 representantes de Lebiasinidae e representantes de outras cinco famílias de Characiformes foram examinados como grupo externo adicionais. A análise filogenética incluindo 208 caracteres de diferentes complexos morfológicos, resultou em seis árvores fundamentais com 800 passos. As topologia obtidas recuperam o monofiletismo de Lebiasininae com base em cinco sinapomorfias (4 exclusivas): margem póstero- dorsal do infra-orbital 3 com uma projeção que exclui parcial- ou totalmente o infra-orbital 4 do anel orbital; lâmina dorsal do mesetmóide em forma de “T”; margem póstero-ventral do coracóide truncada, sem uma distinta projeção direcionada posteriormente; ramo infra-orbital do canal látero- sensorial cefálico se estendendo ao antorbital; mancha na base dos raios médios da nadadeira caudal presente. Por outro lado, estas refutam o monofiletismo de Lebiasina e Piabucina e a sinoníma entre eles é formalmente proposta. O resultado da análise corrobora o monofiletismo de Lebiasinidae (com base em 20 sinapomorfias, 12 exclusivas) e todas as suas demais subunidades. São discutidos eventos graduais de miniaturização em Pyrrhulininae, aparentemente associados a diversos caracteres redutivos, e a evolução dos modos de vida observados em Nannostomus e Pyrrhulinina. Como resultado da revisão taxonômica, 19 das as 22 espécies nominais de Lebiasininae são consideradas válidas e sete são descritas como espécies novas. Entre elas, constam tês novas espécies que até então eram erroneamente classificadas como Lebiasina bimaculata. Todas as espécies são redescritas e novas diagnoses são propostas para cada uma delas. Adicionalmente, uma chave de identificação das espécies (válidas e novas) é apresentada. vii Abstract The subfamily Lebiasininae has been traditionally divided into two genera: Lebiasina and Piabucina. Historically, such division has been contested due to the inconsistency of the diagnostic characters differring both genera. To investigate the relationships among the Lebiasininae and the validity of Lebiasina and Piabucina, 59 representatives of the Lebiasinidae were compared, as well as representatives of five other characiform families as additional outgroups. The phylogenetic analysis, including 208 characters, obtained from different morphological complexes, yielded six fundamental trees with 800 steps. The topologies obtained recover the monophyly of the Lebiasininae based in five synapomorphies (four exclusive): postero-dorsal margin of infraorbital 3 with a projection that excludes patrially or completelly the infraorbital 4 from the orbital rim; dorsal surface of the mesethmoid “T” shaped; postero-ventral margin of coracoid truncate, without a distinct posteriorly projected process; infraorbital ramus of latero-sensory canal extending onto the antorbital; caudal blotch present. On the other hand, the topologies refute the monophyly of Lebiasina and Piabucina, and their synonymy is formally proposed herein. The analysis also recovers the Lebiasinidae as a monophyletic assemblage (based in 20 synapomorhies, 12 exclusive) as well as all other subunities of the family. Gradual miniaturization events apparently associated with several reductive characters and the evolution of life strategies in Nannostomus and Pyrrhulinina are discussed. As a result of the taxonomic review, 19 of the 22 nominal species are considered valid and seven are described as new. Among them, there are three species so far identified as Lebiasina bimaculata. All species are redescribed, their distribution is extended and new diagnoses are proposed to each of them. Additionally, an identification key to the species (valid and new) is presented. viii Agradecimentos Primeiramente gostaria de agradecer à minha família por todo o apoio incondicional dado desde sempre e principalmente durante esses quatro anos de São Paulo. Aos meus filhotinhos por todas as vezes que voltei pra casa e fui recebido com sorrisos contendo pura felicidade por me ver. Ao Marcelo e a Miroca, as pessoas que lá atrás me incentivaram bastante a extrapolar os limites de “casa” e vir fazer o doutorado em outra cidade, outro estado. Apesar das diversas vezes em que me senti um estranho numa terra estranha, valeu cada dia. Muito obrigado pelo empurrão! Durante esses anos de MZUSP tantas pessoas passaram por aqui que acho injusto correr o risco de esquecer alguém na hora listar as pessoas e a importância de cada uma delas no processo, mas tenho certeza de que cada uma delas sabe o papel que teve nessa corrida. Meus agradecimentos especiais vão: para os professores da seção, por toda a ajuda e pelo prazer das histórias contadas na Copa durante o café ou as drinekerias; ao Mario pela orientação, amizade e todas as conversas (sérias, pouco séries ou nada sérias) que tivemos ao longo desses anos; pro Gabiru, pelo companheirismo e amizade ao longo de quase cinco anos lutando contra nossas teses e as forças tentadoras da “deusa da procrastinação”; pra Carinão (mais uma vez) pela amizade, carinho e apoio; pro Pedrinho, pela amizade quase secular e as horas de boteco; pro Flávio, por todas as discussões enriquecedoras (e outras nem tanto, mas de igual valor); pro Cris, também por todas as discussões sobre morfologia de Characiformes; ao Kawada pela ajuda com as automontagens,
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