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Fernando Willyan Trevisan Leivas.Pdf 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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