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Taxonomia E Filogenia Do Gênero Aramides Pucheran, 1845 (Aves: Gruiformes: Rallidae) Taxonomy and Phylogeny of the Genus Aramid

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Taxonomia E Filogenia Do Gênero Aramides Pucheran, 1845 (Aves: Gruiformes: Rallidae) Taxonomy and Phylogeny of the Genus Aramid I Rafael Sobral Marcondes Taxonomia e filogenia do gênero Aramides Pucheran, 1845 (Aves: Gruiformes: Rallidae) Taxonomy and phylogeny of the genus Aramides Pucheran, 1845 (Aves: Gruiformes: Rallidae) São Paulo 2013 II Rafael Sobral Marcondes Taxonomia e filogenia do gênero Aramides Pucheran, 1845 (Aves: Gruiformes: Rallidae) Taxonomy and phylogeny of the genus Aramides Pucheran, 1845 (Aves: Gruiformes: Rallidae) Dissertação apresentada ao Instituto de Biociências da Universidade de São Paulo para a obtenção de Título de Mestre em Ciências, na Área de Zoologia. Orientador: Luís Fábio Silveira São Paulo 2013 III Marcondes, Rafael Sobral Taxonomia e filogenia do gênero Aramides Pucheran, 1845 (Aves: Gruiformes: Rallidae) VIII+254p. Dissertação (Mestrado) - Instituto de Biociências da Universidade de São Paulo. Departamento de Zoologia. 1. Aramides 2. Taxonomia 3. Filogenia I. Universidade de São Paulo. Instituto de Biociências. Departamento de Zoologia. Comissão Julgadora: Prof(a). Dr(a). Prof(a). Dr(a). Prof. Dr. Luís Fábio Silveira Orientador IV Agradecimentos A toda minha família, pelo apoio e confiança em todas as minhas decisões e também por compreender minhas ausências durante as diversas viagens. Ao Luís Fábio Silveira, cuja orientação, confiança e oportunidades ao longo dos últimos anos contribuíram fundamentalmente para minha formação como zoólogo. Aos colegas e amigos da Seção de Aves do Museu de Zoologia da USP, pela agradável convivência diária, pelas discussões, ideias e ensinamentos nos momentos de seriedade, e pela diversão e risadas nos momentos de descontração. A todos que ajudaram em diversos aspectos durante esta pesquisa, seja com os mapas (Marco), com as análises estatísticas (Vagner), com as análises filogenéticas (Guilherme), que desenharam figuras (Glaucia), que revisaram e comentaram rascunhos prévios (Guilherme e Glaucia), ou que simplesmente palpitaram e discutiram comigo sobre as saracuras (muitas pessoas para nomear). Especialmente ao Guilherme Garbino, por ter lido, comentado e discutido versões preliminares de toda a dissertação (alguns trechos mais de uma vez) e principalmente também pelos sete anos de amizade e interesses zoológicos compartilhados. Aos amigos Thiago V. V. Costa e Vagner Cavarzere, pela companhia na viagem aos EUA. O que poderiam ter sido apenas três entediantes semanas estudando espécimes acabou sendo uma viagem extremamente agradável e divertida, além de ter resultado em outros inesperados frutos científicos. Aos amigos Glaucia Del-Rio e Vinicius Tonetti, pela igualmente prazerosa companhia na viagem a Belém para visitar o MPEG. Às instituições visitadas e seus curadores, funcionários e alunos que me receberam: Dr. Alexandre Aleixo, Fátima Lima e Leonardo Moura (MPEG); Dr. Marcos Raposo, Claydson Assis e Daniel Figueira (MNRJ); Ligia M. Abe (MHNCI); Dr. Robert Prys-Jones e Mark Adams (BMNH); Dra. V Sylke Frahnert e Pascal Eckhoff (ZMB); Marie Portas (MNHN); Dr. Joel Cracraft, Paul Sweet e Merle Okada (AMNH); David Willard e Mary Hennan (FMNH). A Leonardo Moura e Cathy Bechtoldt, pela hospedagem em Belém e Chicago, respectivamente. A Vitor Piacentini, Glaucia Del-Rio, Guilherme Garbino, Vagner Cavarzere, Thiago V. V. Costa, Ulf Johansson (Naturhistoriska Riksmuseet, Suécia), Kathy Molina (University of California, EUA), Dominika Mierzwa-Szymkowiak (Muzeum i Instytut Zoologii, Polônia) e René Corado (Western Foundation of Vertebrate Zoology, EUA) por fotografar espécimes depositados em coleções que não pude visitar pessolamente. A todos os colaboradores que gentilmente me enviaram gravações de seus acervos pessoais, em especial aqueles que contribuíram com as raras gravações de Aramides “cajaneus” da América Central: Jesse Fagan, John Moore, Luis E. Vargas e Luis Sandoval (Laboratorio de Bioacustica, Universidad de Costa Rica). À Fonoteca Neotropical Jacques Vieilliard da UNICAMP e à Macaulay Library da Cornell University (especialmente Matthew Meddler, Greg Budney e Tammy Bishop), por me enviar gravações lá depositadas. Ao Instituto de Biociências e ao Museu de Zoologia da USP, seus funcionários, professores e colaboradores, por terem fornecido a infraestrutura e o apoio que tornaram esse trabalho possível. Ao Prof. Dr. Marcelo Rodrigues de Carvalho, pelo apoio como coordenador do Programa de Pós-Graduação em Zoologia e por ter me escrito cartas de recomendação em duas ocasiões diferentes. Ao CNPq, pela bolsa concedida (processo número 130272/2011-6). Ao American Museum of Natural History, pelo auxílio financeiro (Collection Study Grant Award) concedido para minha visita. VI Ao meu tio Arthur Marcondes, pela passagem aérea que permitiu minhas visitas às coleções européias. A todos os naturalistas e cientistas, de diversas gerações, que coletaram e gravaram os espécimes que foram a fundação deste trabalho. Desde Johann Natterer e Alfred Russel Wallace, passando por Helmut Sick e Ted Parker, até meus próprios colegas e amigos de laboratório, esta dissertação é resultado também do esforço deles. VII Índice Introdução Geral........................................................................................................... 1 Capítulo 1. Revisão taxonômica e distribuição do gênero Aramides Pucheran, 1845, com base em caracteres morfológicos externos e vocais.............................................. 11 1.1 Introdução......................................................................................................... 14 1.2. Material e Métodos......................................................................................... 20 1.2.1. Morfologia............................................................................................. 20 1.2.1.1. Espécimes estudados.................................................................. 20 1.2.1.2. Caracteres de plumagem............................................................ 21 1.2.1.3. Caracteres morfométricos e análises estatísticas....................... 21 1.2.2. Vocalizações.......................................................................................... 24 1.2.3. Distribuição geográfica.......................................................................... 28 1.2.4. Critérios para delimitação de espécies.................................................. 28 1.3. Resultados........................................................................................................ 30 1.3.1. Variação morfológica e vocal e limites específicos............................... 30 1.3.1.1. Variação de plumagem............................................................... 30 1.3.1.1.1. Aramides ypecaha............................................................ 31 1.3.1.1.2. Aramides wolfi................................................................. 32 1.3.1.1.3. Aramides mangle............................................................. 33 1.3.1.1.4. Aramides cajaneus........................................................... 33 1.3.1.2. Variação vocal............................................................................. 41 1.3.1.4. Morfometria............................................................................... 42 1.3.2. Sinopse do gênero Aramides................................................................. 44 1.4. Discussão.......................................................................................................... 74 Capítulo 2. Análise filogenética do gênero Aramides Pucheran, 1845, com base em caracteres tegumentares e vocais................................................................................. 91 2.1. Introdução........................................................................................................ 94 2.1.1. Histórico filogenético do gênero Aramides........................................... 94 2.1.2. Uso de caracteres tegumentares e vocais para reconstruções filogenéticas em aves ..................................................................................... 96 2.1.3. Objetivo................................................................................................. 98 2.2. Material e Métodos......................................................................................... 98 2.2.1. Amostragem taxonômica e material analisado..................................... 98 2.2.2. Delimitação dos caracteres................................................................... 101 2.2.2.1. Caracteres tegumentares........................................................... 101 VIII 2.2.2.2. Caracteres vocais........................................................................ 103 2.2.3 Análise filogenética................................................................................ 104 2.3. Resultados........................................................................................................ 106 2.3.1. Caracteres............................................................................................. 106 2.3.2. Análises filogenéticas............................................................................ 119 2.3.2.1. Análise A..................................................................................... 119 2.3.2.2. Análise B..................................................................................... 120 2.3.2.3. Análise C..................................................................................... 121 2.3.2.4. Análise D..................................................................................... 121 2.3.2.5. Análise E.....................................................................................
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