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Suzete Rodrigues Gomes SUZETE RODRIGUES GOMES Filogenia morfológica de Veronicellidae, filogenia molecular de Phyllocaulis Colosi e descrição de uma nova espécie para a família (Mollusca, Gastropoda, Pulmonata). 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 Doutor em Biologia Animal. Área de Concentração: Biologia Comparada Orientador: Dra. Inga Veitenheimer Mendes Co-orientador: Dr. José Willibaldo Thomé UNIVERSIDADE FEDERAL DO RIO GRANDE DO SUL Porto Alegre 2007 Filogenia morfológica de Veronicellidae, filogenia molecular de Phyllocaulis Colosi e descrição de uma nova espécie para a família (Mollusca, Gastropoda, Pulmonata) SUZETE RODRIGUES GOMES Aprovada em ________/_________/________ Banca examinadora ________________________________________________________________ Dra. Norma Campos Salgado ________________________________________________________________ Dr. Gervásio Silva Carvalho _______________________________________________________________ Dra. Suzana Bencke Amato ii SUMÁRIO AGRADECIMENTOS............................................................................... vii RESUMO................................................................................................. viii CAPÍTULO 1 ASPECTOS GERAIS DE VERONICELLIDAE........................................ 1 Morfologia, Sistemática e Importância de Veronicellidae........................ 2 Principais Estudos, Breve Histórico e Distribuição dos Gêneros............... 4 Região Etiópica............................................................................ 4 Filicaulis.................................................................................... 5 Laevicaulis................................................................................. 6 Pseudoveronicella....................................................................... 6 Região Australiana e Oriental..................................................... 8 Filicaulis.................................................................................... 9 Laevicaulis................................................................................. 9 Sarasinula................................................................................. 10 Semperula................................................................................. 10 Valiguna.................................................................................... 11 Veronicella................................................................................. 12 Região Neotropical..................................................................... 12 Angustipes................................................................................. 13 Belocaulus................................................................................. 14 Colosius..................................................................................... 15 Diplosolenodes........................................................................... 15 Forcartulus................................................................................ 16 Heterovaginina........................................................................... 16 Latipes...................................................................................... 17 Leidyula.................................................................................... 18 Microveronicella.......................................................................... 19 Montivaginulus........................................................................... 19 Novovaginula............................................................................. 20 Phyllocaulis................................................................................ 20 Potamojanuarius......................................................................... 21 Sarasinula................................................................................. 22 Simrothula................................................................................. 22 Vaginulus.................................................................................. 23 Veronicella.....................….......................................................... 24 iii Zilchulus.................................................................................... 24 Análise do Atual Conhecimento sobre Veronicellidae e Objetivos da Tese........................................................................................................ 25 Referências Bibliográficas ...................................................................... 26 CAPÍTULO 2 A ORIGEM GONDUÂNICA DE VERONICELLIDAE (GASTROPODA, SOLEOLIFERA): EVIDÊNCIAS A PARTIR DE UMA ANÁLISE CLADÍSTICA BASEADA EM CARACTERES MORFOLÓGICOS.................................... 33 Resumo....................................................................................... 34 Introdução.................................................................................. 35 Material e Métodos...................................................................... 37 Discussão dos Caracteres............................................................ 39 Resultados.................................................................................. 59 Discussão.................................................................................... 63 Agradecimentos.......................................................................... 67 Referências................................................................................. 67 Tabelas........................................................................................ 74 Figuras........................................................................................ 76 CAPÍTULO 3 MOLECULAR PHYLOGENY OF THE LAND SLUGS OF THE GENUS PHYLLOCAULIS (MOLLUSCA; SOLEOLIFERA; VERONICELLIDAE) EVALUATED IN LIGHT OF MORPHOLOGICAL AND GEOGRAPHICAL EVIDENCES.............................................................................. 125 Abstract...................................................................................... 127 Introduction................................................................................ 128 Material and Methods.................................................................. 129 Results........................................................................................ 132 Discussion.................................................................................. 133 Conclusions................................................................................. 136 Acknowledgments....................................................................... 136 References................................................................................. 137 Tables......................................................................................... 141 Figures........................................................................................ 143 iv CAPÍTULO 4 A NEW SPECIES OF SIMROTHULA (GASTROPODA, SOLEOLIFERA, VERONICELLIDAE) FROM NORTHERN BRAZIL………………………………….…… 148 Abstract...................................................................................... 149 Introduction................................................................................ 149 Material and Methods.................................................................. 150 Systematics................................................................................. 151 External characteristics............................................................... 151 Digestive system......................................................................... 152 Pedal and pallial nerves, pedal aortic system.............................. 152 Pedal gland................................................................................. 153 Reproductive system................................................................... 153 Discussion................................................................................... 153 Acknowledgments....................................................................... 154 Literature cited............................................................................ 155 Figures........................................................................................ 157 CAPÍTULO 5 CONSIDERAÇÕES FINAIS............................................................ 162 ANEXOS................................................................................. 166 Normas: Zoological Journal of the Linnean Society 167 .................... Normas: Molecular Phylogenetics and Evolution 170 ........................ Normas: Zootaxa................................................................... 172 v DEDICATÓRIA Ao meu mestre Professor José Willibaldo Thomé, por todos os ensinamentos vi AGRADECIMENTOS Ao Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), pelo apoio financeiro concedido no período de março de 2003 a fevereiro de 2007, sem o qual eu não poderia ter realizado este trabalho. Aos meus orientadores: Profa. Inga, por ter acreditado em mim e pela orientação e amizade durante os quatro anos nos quais o trabalho foi desenvolvido; ao Prof. José Willibaldo por ter sido meu mestre não só durante este período, mas durante os últimos 11 anos da minha vida. A ele agradeço pelos conhecimentos transmitidos, não só sobre Veronicellidae, estudada por ele a mais de 40 anos, mas também por seus sempre tão sábios ensinamentos. Ao seu lado
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