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Mollusca, Gastropoda) Baseada Em Morfologia Comparada

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Mollusca, Gastropoda) Baseada Em Morfologia Comparada Paulino José Soares de Souza Jr. Análise cladística da família Marginellidae (Mollusca, Gastropoda) baseada em morfologia comparada. São Paulo 2007 i Paulino José Soares de Souza Jr. Análise cladística da família Marginellidae (Mollusca, Gastropoda) baseada em morfologia comparada. 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(a): Luiz R. L. de Simone São Paulo 2007 ii Ficha Catalográfica Souza, Paulino José Soares, Jr. Análise cladística da família Marginellidae (Mollusca, Gastropoda) baseada em morfologia comparada x+129 pp. Tese (Doutorado) - Instituto de Biociências da Universidade de São Paulo. Departamento de Zoologia. 1. Palavra- Mollusca 2. Análise cladística 3. Anatomia I. Universidade de São Paulo. Instituto de Biociências. Departamento de Zoologia. Comissão Julgadora: ________________________ _____ _______________________ Prof(a). Dr(a). Prof(a). Dr(a). _________________________ ____________________________ Prof(a). Dr(a). Prof(a). Dr(a). Prof. Dr. Luiz R. L. de Simone Orientador iii Dedico esta tese à memória do meu tio, Almirante José Gerardo Theófilo Albano de Aratanha, que me iniciou no mundo dos moluscos. iv “Da natureza, existe em nossas mentes, um desejo insaciável de conhecer a verdade” Cícero, Tusculanarum Disputationum:1.2.44 v Sumário SUMÁRIO...................................................................................................................................................... V AGRADECIMENTOS.................................................................................................................................VI RESUMO ...................................................................................................................................................VIII ABSTRACT ................................................................................................................................................... X 1. INTRODUÇÃO .......................................................................................................................................... 1 1.1. GENERALIDADES .................................................................................................................................... 1 1.2. PALEONTOLOGIA .................................................................................................................................... 3 1.3. HISTÓRICO TAXONÔMICO ....................................................................................................................... 4 1.3.1. Posição dos Marginellidae em Gastropoda ................................................................................... 4 1.3.2. Taxonomia supraespecífica dos Marginellidae..............................................................................6 1.4. JUSTIFICATIVA PARA O PRESENTE ESTUDO ........................................................................................... 16 1.5. OBJETIVOS............................................................................................................................................ 18 2. MATERIAL E MÉTODOS ..................................................................................................................... 19 2.1. ABREVIATURAS:................................................................................................................................... 19 2.1.1 - Coleções:..................................................................................................................................... 19 2.1.2 - Anatomia:.................................................................................................................................... 19 2.2. MATERIAL EXAMINADO:...................................................................................................................... 21 2.2.1. Cystiscidae.................................................................................................................................... 21 2.2.2. Volutidae ...................................................................................................................................... 21 2.2.3. Marginellidae ............................................................................................................................... 22 2.3. MÉTODOS ............................................................................................................................................. 29 2.3.1. Anatomia ...................................................................................................................................... 29 2.3.2 Ilustrações ..................................................................................................................................... 30 2.3.3. Análise Cladística......................................................................................................................... 30 2.3.4. Otimização: .................................................................................................................................. 32 2.3.5. Programas de computador auxiliares na análise cladística: ....................................................... 32 3. RESULTADOS......................................................................................................................................... 33 3.1. CARACTERES........................................................................................................................................ 33 3.1.1. Concha.......................................................................................................................................... 33 3.1.2. Massa céfalo-pediosa ................................................................................................................... 42 3.1.3. Órgãos do manto e sistemas circulatório e excretor.................................................................... 48 3.1.4. Sistema digestório ........................................................................................................................ 56 3.1.4. Sistema reprodutor ....................................................................................................................... 91 3.2. MATRIZ DE CARCTERES ........................................................................................................................ 99 3.3. CLADOGRAMA.................................................................................................................................... 100 4. DISCUSSÃO ........................................................................................................................................... 103 5. CONCLUSÕES ...................................................................................................................................... 116 6. REFERÊNCIAS BIBLIOGRÁFICAS.................................................................................................. 117 vi Agradecimentos Meu caminho até a conclusão desta tese foi longo, e não teria chegado ao fim dele não fora o auxílio de meus orientadores: Dr. Nélson Papavero, que aceitou me orientar no início do projeto; Prof. Dr. Mario C. C. de Pinna, por ter dado a chance de reiniciar o caminho, quando eu não acreditava mais que ainda era possível; e Prof. Dr. Luiz Ricardo L. Simone, que além de ter me ensinado tudo que sei sobre anatomia de moluscos, sempre me brindou com sua amizade e apoio, e cuja obstinação e zelo científicos, só posso aspirar. Também, gostaria de agradecer ao Dr. José Luiz de Moreira Leme, por me receber na Seção de Malacologia do Museu de Zoologia da USP. Sou muito grato ao Dr. Gary A. Coovert e à sua esposa Holly K. Coovert, por me receberam pra uma agradabilíssima e produtiva estada em sua residência e por me cederam, além de várias cópias de artigos consultados, originais de toda a série Marginella Marginallia. Às bibliotecárias do MZUSP, Teresa Beatriz Nunes Guimarães, Cláudia Alves de Melo, Janete da Rocha, Carmina Lupo, meus por todo auxílio bibliográfico, sempre prestado com boa vontade e simpatia, tão raras nos dias de hoje. E um agradecimento muito especial à Dione Seripierri, pela minuciosa revisão da lista bibliográfica. Minha longa estada em São Paulo teria sido muito mais solitária sem meus amigos e colegas do MZUSP, em especial Carlo Magenta da Cunha (que também me conseguiu lotes de Eratoidea scalaris), Cibele Bragagnolo, Gilson Iak Ximenes, Carlos Augusto Figueiredo e Flávio C. C. de Lima. Devo um agradecimento especial a Cristiano Moreira que sempre me auxiliou com meus minhas dúvidas em relação aos softwares de análise filogenética, mesmo quando seu próprio tempo era escasso. Ao amigo e co-fundador do CML Eric P. Gonçalves, sou profundamente grato por participar em, praticamente, todos vii os aspectos desta tese, até o último minuto! Agradeço, especialmente, ao meu amigo Antônio Cornetta (Black Hawk Computers), não só pela companhia como também pela consultoria em assuntos de informática. A José Coltro, minha sincera gratidão, por facilitar minha na coleta de material no sul da Bahia, onde coletei os exemplares de Prunum fulminatum e por ter cedido o tão precioso lote de Marginellona gigas estudado aqui, um espécime raro que ajudou muito nas conclusões desta tese. Aos curadores que cederam a maior parte dos espécimes aqui estudados: Dr. Miroslaw G. Harasewych (Smithsonian Museum), Dr. Winston Ponder e Sr. Ian Locke, (Australian Museum), Dra. Nancy Voss (Marine
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