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Revisão Taxonômica E Análise Filogenética De Paraphasma Redtenbacher, 1906 (Phasmatodea: Pseudophasmatidae: Stratocleinae)

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Revisão Taxonômica E Análise Filogenética De Paraphasma Redtenbacher, 1906 (Phasmatodea: Pseudophasmatidae: Stratocleinae) UNIVERSIDADE DE SÃO PAULO MUSEU DE ZOOLOGIA Pedro Ivo Chiquetto Machado Revisão taxonômica e análise filogenética de Paraphasma Redtenbacher, 1906 (Phasmatodea: Pseudophasmatidae: Stratocleinae) Taxonomic revision and phylogenetic analysis of Paraphasma Redtenbacher, 1906 (Phasmatodea: Pseudophasmatidae: Stratocleinae) Volume I São Paulo 2019 Pedro Ivo Chiquetto Machado Revisão taxonômica e análise filogenética de Paraphasma Redtenbacher, 1906 (Phasmatodea: Pseudophasmatidae: Stratocleinae) Taxonomic revision and phylogenetic analysis of Paraphasma Redtenbacher, 1906 (Phasmatodea: Pseudophasmatidae: Stratocleinae) Volume I Versão Original Tese apresentada ao Programa de Pós- Graduação do Museu de Zoologia da Universidade de São Paulo para obtenção do título de Doutor em Ciências, na Área de Sistemática, Taxonomia Animal e Biodiversidade. Orientadora: Profa. Dra. Eliana Marques Cancello São Paulo 2019 Autorizo a reprodução e divulgação total ou parcial deste trabalho, por qualquer meio convencional ou eletrônico, para fins de estudo e pesquisa, desde que citada a fonte. I authorize the reproduction and dissemination of this work in part or entirely by any electronic or conventional means, for study and research, provided the source is cited. Catalogação na Publicação Serviço de Biblioteca e Documentação Museu de Zoologia da Universidade de São Paulo Machado, Pedro Ivo Chiquetto Revisão taxonômica e análise filogenética de Paraphasma Redtenbacher, 1906 (Phasmatodea: Pseudophasmatidae: Stratocleinae) = Taxonomic revision and phylogenetic analysis of Paraphasma Redtenbacher, 1906 (Phasmatodea: Pseudophasmatidae: Stratocleinae) – Pedro Ivo Chiquetto Machado; orientadora Eliana Marques Cancello. São Paulo, 2019. 2v. Tese apresentada ao Programa de Pós-Graduação do Museu de Zoologia da Universidade de São Paulo para a obtenção do título de Doutor em Ciências, na área de Sistemática, Taxonomia Animal e Biodiversidade. Versão original 1. Phasmatodea – análise filogenética. 2. Stratocleinae. 3. Pseudophasmatidae. I. Cancello, Eliana Marques, orient. II. Título. CDU 595.72 CRB 8 /3805 MACHADO, Pedro Ivo Chiquetto Revisão taxonômica e análise filogenética de Paraphasma Redtenbacher, 1906 (Phasmatodea: Pseudophasmatidae: Stratocleinae) Taxonomic revision and phylogenetic analysis of Paraphasma Redtenbacher, 1906 (Phasmatodea: Pseudophasmatidae: Stratocleinae) Tese apresentada ao Programa de Pós-Graduação do Museu de Zoologia da Universidade de São Paulo para obtenção do título de Doutor em Ciências, na Área de Sistemática, Taxonomia Animal e Biodiversidade. Aprovado em:______/______/________ Banca examinadora Prof. Dr. _________________________________ Instituição: ___________________________ Julgamento: ______________________________ Assinatura:___________________________ Prof. Dr. _________________________________ Instituição: ___________________________ Julgamento: ______________________________ Assinatura:___________________________ Prof. Dr. _________________________________ Instituição: ___________________________ Julgamento: ______________________________ Assinatura:___________________________ Prof. Dr. _________________________________ Instituição: ___________________________ Julgamento: ______________________________ Assinatura:___________________________ Prof. Dr. _________________________________ Instituição: ___________________________ Julgamento: ______________________________ Assinatura:___________________________ AGRADECIMENTOS À minha orientadora, Profa. Eliana Marques Cancello, pelas importantes sugestões, críticas e discussões, e por toda a dedicação em me orientar no estudo de um grupo diferente de sua especialidade. Aos curadores de instituições estrangeiras e brasileiras que emprestaram material para este estudo ou que me receberam em visitas às coleções: Dra. Susanne Randolf e Harald Bruckner (Naturhistorisches Museum Wien); Dr. Ben Price (Natural History Museum); Amoret Spooner (Oxford University Museum of Natural History); Dr. Roberto Zucchi (ESALQ-USP); Dr. Marcio Oliveira (INPA); Dr. Miguel Monné (MN-UFRJ); Dr. Orlando Silveira (MPEG); Dra. Luciane Marinoni (UFPR); Dr. Paulo Motta (UnB); Dr. Freddy Bravo (UEFS); Dr. Fernando Vaz de Mello (UFMT); Dr. Ricardo Andreazze (in memoriam) (UFRN); Dra. Jane Costa (FIOCRUZ). À Profa. Adriana Coletto Morales, por sua importante colaboração no projeto, tendo me recebido no Laboratório de Biologia Evolutiva da Universidade Estadual Paulista “Júlio de Mesquita Filho” (UNESP – Campus de Jaboticabal) e disponibilizado a infraestrutura do laboratório para este trabalho, além de ter e me orientado e auxiliado nos procedimentos para obtenção de sequências genéticas; também a Amanda de Faria, pela disponibilidade em me auxiliar durante os dias em que estive na UNESP. A Oskar Conle, pelas sugestões, discussões taxonômicas e fornecimento de imagens para este estudo. Ao Prof. Marcelo Duarte e a Renato Silva, pelo empréstimo de equipamentos de fotografia utilizados para a obtenção de diversas imagens desta tese. A Ana Vasques e Carlos Campaner, que me auxiliaram em diversas situações no dia a dia de trabalho no MZUSP. À equipe da biblioteca do MZUSP, pelo auxílio na busca de obras importantes para a realização deste trabalho e, em especial, a Dione Seripierri, por sanar diversas dúvidas relacionadas à citação de artigos e à formatação de referências bibliográficas. A Fernando Domenico, por ter me ensinado técnicas de dissecção que foram de grande importância para a realização deste estudo. A Maurício Rocha e Juan Pablo Botero, pelo auxílio na realização de análises filogenéticas. A Joice Constantini, Nara Barbosa, Tiago Carrijo, Rafaella Santos e Carolina Cuezzo, pela colaboração e companhia em expedições de coleta em busca de bichos-paus e cupins. A Raphael Heleodoro, pelo fornecimento de imagens para este estudo. Aos amigos do Laboratório de Ortopteroides e todos os amigos do Museu, pela amizade e ótimos momentos compartilhados durante esses quatro anos. Aos amigos do Projeto Phasma – Alvaro, Edgar, Phillip e Victor – pela amizade e tantos conhecimentos trocados a respeito dos fantásticos bichos-paus. Aos meus pais e minhas irmãs, por todo o apoio, amizade e convivência, tendo sempre demonstrado interesse pelo meu trabalho. À minha esposa, Alessandra, pelo auxílio e companhia em muitas expedições de coleta deste estudo, pelas diversas sugestões e críticas ao texto, e por continuar sendo, após mais de dez anos juntos, a maior amizade e maior certeza da minha vida. À Fundação de Amparo à Pesquisa do Estado de São Paulo, pelo apoio financeiro a este trabalho através da concessão de Bolsa de Doutorado (Processo FAPESP 2015/08808-2) e Bolsa Estágio de Pesquisa no Exterior (Processo FAPESP 2017/08022-4). À Coordenação de Aperfeiçoamento de Pessoal de Nível Superior, pelo apoio financeiro no início do projeto através da concessão de bolsa CAPES/Demanda Social (nível: doutorado). Eu te espero chegar vendo os bichos, sozinho, na noite. Distração de quem quer esquecer o seu próprio destino. O Terço, Criaturas da Noite RESUMO A taxonomia de Phasmatodea baseia-se essencialmente em caracteres de morfologia externa e características do ovo. A genitália masculina, estrutura com grande relevância taxonômica em outras ordens de insetos, foi examinada em poucas espécies de bicho-pau, sendo a sua morfologia ainda muito pobremente explorada para fins taxonômicos e filogenéticos. Das poucas análises filogenéticas realizadas até hoje em Phasmatodea, a maioria foi baseada em dados moleculares e focou nas relações entre grandes grupos, com apenas dois estudos abordando relações intragenéricas. Paraphasma Redtenbacher, 1906 é um gênero neotropical de bichos-paus pequenos e de aspecto delicado, com ambos os sexos possuindo asas posteriores bem desenvolvidas. Incluía, até o início deste estudo, dez espécies válidas, oito das quais com registro para o Brasil; trata-se de um gênero relativamente bem representado nas coleções brasileiras de Phasmatodea. Porém, assim como grande parte dos gêneros brasileiros de bichos-paus, Paraphasma encontrava-se taxonomicamente mal definido, e o conhecimento sobre suas espécies era praticamente restrito às descrições originais, normalmente muito superficiais, e a informações vagas de distribuição geográfica. O objetivo deste estudo foi realizar uma revisão taxonômica e análise filogenética de Paraphasma, buscando explorar o potencial taxonômico da genitália masculina de Phasmatodea. Foram examinados 407 exemplares (incluindo 38 tipos), estando representadas todas as espécies de Paraphasma e 18 espécies como grupo-externo da análise filogenética. A genitália masculina foi examinada e ilustrada para 22 espécies; foram verificadas prováveis homologias entre as espécies e propostos nomes para as regiões e estruturas da genitália. Foram definidos 80 caracteres morfológicos para a análise filogenética, dos quais 32 referem- se à genitália masculina. Como resultado, Paraphasma passa a ter uma configuração bastante distinta da inicial, incluindo nove espécies, das quais três são espécies novas identificadas durante o estudo. O gênero foi redescrito, com base em morfologia externa e em características da genitália masculina e do ovo. Foram descritas as três espécies novas e redescritas as demais espécies de Paraphasma, incluindo a primeira descrição do macho de Paraphasma minus, do ovo de seis espécies e da genitália masculina de oito. Foram propostos dois novos gêneros (chamados aqui pelos nomes fictícios “Aus” e “Bus”) para duas espécies previamente em Paraphasma, resultando nas
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