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Fernando Willyan Trevisan Leivas Análise Cladística de Omalodini Kryzhanovskij, 1972 (Coleoptera, Histeridae, Histerinae) Tese apresentada à Coordenação do Programa de Pós-Graduação em Ciências Biológicas, Área de concentração em Entomologia, da Universidade Federal do Paraná, como requisito parcial para obtenção do título de Doutor em Ciências Biológicas. Orientadora: Profa. Dra. Lúcia Massutti de Almeida Co-orientadora: Profa. Dra. Carla de Lima Bicho Curitiba - Paraná 2012 Dedico esta pesquisa à minha família e minhas orientadoras Dras. Lúcia Massutti de Almeida e Carla de Lima Bicho. Agradeço a Deus pela maneira que tem conduzido meus caminhos. "...the past constrains the future only to the extent that it has structured the present... Hull (1988)". ii AGRADECIMENTOS Nestas palavras, agradeço a todos que contribuíram de alguma maneira para a realização deste trabalho. A Universidade Federal do Paraná, ao Programa de Pós-graduação em Entomologia, e ao Laboratório de Sistemática e Bioecologia de Coleoptera (Insecta) pela infraestrutura. Ao CNPq pela concessão da bolsa de estudos. Ao Programa Nacional de Cooperação Acadêmica (PROCAD) pelo auxílio financeiro durante o intercâmbio com o Instituto Nacional de Pesquisas da Amazônia (INPA). Aos docentes do Programa de Pós-graduação em Entomologia, da Universidade Federal do Paraná, pela transferência de conhecimento. Aos curadores das Coleções Entomológicas, nacionais e estrangeiras, e colecionadores particulares pelo empréstimo de material que viabilizou este estudo, em especial ao Dr. Nicolas Degallier, pelo auxílio nos estudos com Histeridae. Ao Drs. Nicolas Degallier, Michael S. Caterino e Alexey K. Tishechkin e Msc. Daniel de Pessoa Moura pelas ricas discussões sobre os resultados preliminares e finais da tese. A Willey Hening Society e autores do TNT pelo livre acesso do programa (Goloboff, P.A., Farris, J.S. & Nixon, K.C. (2008) TNT, a free program for phylogenetic analysis.Cladistics, 24, 774-786). Ao Centro de Microscopia Eletrônica e Programa Taxon line, da Universidade Federal do Paraná, pelo trabalho com as imagens de microscópio eletrônico e coloridas, respectivamente. Assim também a todos os funcionários da Universidade Federal do Paraná, PRPPG e Departamento de Zoloogia, pelo auxílio nas questões requisitadas. Aos Drs. Mirna Martins Casagrande, Olaf Herman Hendrik Mielke, Paschoal Coelho Grossi, Eduardo Carneiro e aos MScs. Diego Dolibana, Fernando Maia e Claudivan Matos Maia que colaboraram na coleta de material. Ao Paschoal Coelho Grossi, Everardo Grossi e Lenora Paes Coelho Grossi um agradecimento especial pelo carinho em me hospedar em sua casa durante a visita do Dr. Nicolas Degallier, ao Rio de Janeiro. Aos amigos discentes do Programa de Pós-graduação em Entomologia, assim como aos amigos de longa data, pelos momentos que passamos juntos, com grande alegria, sempre proporcionando trocas culturais. iii A Dra. Cibele Stramare Ribeiro-Costa, todos os colegas e amigos do Laboratório de Sistemática e Bioecologia de Coleoptera, pelas discussões científicas, convívio e colaboração nas pesquisas na tese e extratese. iv SUMÁRIO INTRODUÇÃO ............................................................................................................... 15 Histórico de Omalodini ........................................................................................ 19 MATERIAL E MÉTODOS .............................................................................................. 23 Estudo morfológico .............................................................................................. 23 Material examinado .............................................................................................. 24 Terminologia morfológica .................................................................................... 26 Táxons terminais .................................................................................................. 26 Construção e levantamento dos caracteres ............................................................ 30 Metodologia Cladística ......................................................................................... 31 RESULTADOS E DISCUSSÃO ...................................................................................... 32 Caracteres e matriz final ....................................................................................... 32 Lista dos caracteres utilizados na análise cladística de Omalodini ................. 41 Caracteres não incluídos na análise ............................................................... 51 Resultados da Análise Cladística .......................................................................... 53 Mapeamento das transformações comuns ............................................................. 55 Discussão das Hipóteses Geradas ......................................................................... 62 Monofilia de Omalodini e de suas linhagens (clado 81)................................. 62 Grupo imão de Omalodini ............................................................................. 78 Grupos considerados externos a priori e a posteriori da análise .................... 81 Omalodini: Sistemática Revisada.......................................................................... 88 Classificação estabelecida para Omalodini .................................................... 88 Chave de identificação para os gêneros de Omalodini Kryzhanovskij, 1972 .. 90 CONSIDERAÇÕES FINAIS ........................................................................................... 90 REFERÊNCIAS .............................................................................................................. 91 Figuras ............................................................................................................... 100 Apêndices........................................................................................................... 208 LISTA DE TABELAS Tabela I. Terminais incluídos na análise cladística de Omalodini Kryzhanovskij, 1972 (Coleoptera, Histeridae). .................................................................................................. 28 Tabela II. Matriz de estados dos caracteres utilizados na análise cladística de Omalodini Kryzhanovskij, 1972 (Coleoptera, Histeridae). Dados não observados (?) e dados não aplicáveis (-). ................................................................................................................... 33 Tabela III. Sumário das buscas com diferentes esquemas de pesagem. N= análise normal, CSN= consenso estrito da análise normal, AM= Análise “For monophyly”, CSAM= consenso estrito da análise “For monophyly”, Ck= árvore de consenso estrito, CSP= consenso estrito de todas as árvores geradas pela análise de pesagem implícita perante os distintos valores de k, AR= número de árvores obtidas, L= comprimento das árvores, IC= índice de consistência, IR: índice de retenção, CLF= classificação sensu Mazur (2011), NM= não monofilético, M= Monofilético. ....................................................................... 61 v LISTA DE FIGURAS Figura 1. Hipóteses de relacionamento filogenético existentes para Histerinae. A. Cladograma elaborado com base na proposta de relacionamento entre alguns gêneros de Histerinae segundo Ślipiński e Mazur (1999). B. Cladograma elaborado com base nas propostas de relacionamento entre alguns gêneros de Histerinae segundo Caterino e Vogler (2002). ............................................................................................................................. 17 Figura 2. Terminologia, Histeridae. A. corpo vista dorsal, B. corpo vista ventral e C. pronoto e élitros em vista lateral. .................................................................................... 100 Figura 3. Terminologia, Histeridae. A. cabeça vista ventral; B. cabeça vista dorsal; C. clava antenal; D. maxila; E. perna anterior em face posterior; F. perna anterior em face anterior. ...................................................................................................................................... 101 Figura 4. Terminologia, Histeridae. Macho. A. edeago em vista dorsal; B. edeago em vista ventral; C. oitavo tergito; D. oitavo esternito; E. nono esternito/tergito e décimo tergito. Fêmea. F. bursa copulatríx, oviduto comum, espermateca e glândula espermática; G. coxitos e estilos. ............................................................................................................. 102 Figura 5. Regiões utilizadas para as mensurações das estruturas de Histeridae: A. corpo em vista dorsal, B. corpo em vista ventral, C. edeago e D. fronte. (Cp) Comprimento pronotal; (Ce) comprimento elitral; (Clp) comprimento do lobo prosternal; (Cqp) comprimento da quilha prosternal; (Cpr) comprimento posternal; (Cm) comprimento mesosternal; (Cpe) comprimento do primeiro esterno abdominal visível; (Cte) comprimento do terceiro esterno abdominal visível; (Cpb) comprimento da peça basal; (Cpa) comprimento dos parâmeros; (Clm) comprimento do lobo médio; (Fme) menor distância entre os olhos; (Fma) maior distância entre os olhos. ............................................................................. 103 Figura 6. Aspecto dorsal dos terminais utilizados na análise, grupo externo. A. Epierus sp.; B. Haeterius ferrugineus (Olivier, 1789); C. Alloiodites plaumanni (Reichensperger, 1939); D. Exosternus terminalis (Schmidt, 1889); E. Operclipygus hospes Lewis, 1902; F. Hister putridus Erichson, 1834. Escala 1 mm.
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    Comitato Scientifico per la Fauna d’Italia CHECKLIST AND DISTRIBUTION OF THE ITALIAN FAUNA FAUNA THE ITALIAN AND DISTRIBUTION OF CHECKLIST 10,000 terrestrial and inland water species and inland water 10,000 terrestrial CHECKLIST AND DISTRIBUTION OF THE ITALIAN FAUNA 10,000 terrestrial and inland water species ISBNISBN 88-89230-09-688-89230- 09- 6 Ministero dell’Ambiente 9 778888988889 230091230091 e della Tutela del Territorio e del Mare CH © Copyright 2006 - Comune di Verona ISSN 0392-0097 ISBN 88-89230-09-6 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, without the prior permission in writing of the publishers and of the Authors. Direttore Responsabile Alessandra Aspes CHECKLIST AND DISTRIBUTION OF THE ITALIAN FAUNA 10,000 terrestrial and inland water species Memorie del Museo Civico di Storia Naturale di Verona - 2. Serie Sezione Scienze della Vita 17 - 2006 PROMOTING AGENCIES Italian Ministry for Environment and Territory and Sea, Nature Protection Directorate Civic Museum of Natural History of Verona Scientifi c Committee for the Fauna of Italy Calabria University, Department of Ecology EDITORIAL BOARD Aldo Cosentino Alessandro La Posta Augusto Vigna Taglianti Alessandra Aspes Leonardo Latella SCIENTIFIC BOARD Marco Bologna Pietro Brandmayr Eugenio Dupré Alessandro La Posta Leonardo Latella Alessandro Minelli Sandro Ruffo Fabio Stoch Augusto Vigna Taglianti Marzio Zapparoli EDITORS Sandro Ruffo Fabio Stoch DESIGN Riccardo Ricci LAYOUT Riccardo Ricci Zeno Guarienti EDITORIAL ASSISTANT Elisa Giacometti TRANSLATORS Maria Cristina Bruno (1-72, 239-307) Daniel Whitmore (73-238) VOLUME CITATION: Ruffo S., Stoch F.