Hemiptera: Heteroptera: Pentatomoidea

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Hemiptera: Heteroptera: Pentatomoidea VIVIANA CAUDURO MATESCO SISTEMÁTICA DE THYREOCORIDAE AMYOT & SERVILLE (HEMIPTERA: HETEROPTERA: PENTATOMOIDEA): REVISÃO DE ALKINDUS DISTANT, MORFOLOGIA DO OVO DE DUAS ESPÉCIES DE GALGUPHA AMYOT & SERVILLE E ANÁLISE CLADÍSTICA DE CORIMELAENA WHITE, COM CONSIDERAÇÕES SOBRE A FILOGENIA DE THYREOCORIDAE, E MORFOLOGIA DO OVO DE 16 ESPÉCIES DE PENTATOMIDAE COMO EXEMPLO DO USO DE CARACTERES DE IMATUROS EM FILOGENIAS Tese apresentada ao Programa de Pós-Graduação em Biologia Animal, Instituto de Biociências, Universidade Federal do Rio Grande do Sul, como requisito parcial à obtenção do Título de Doutor em Biologia Animal. Área de concentração: Biologia Comparada Orientadora: Profa. Dra. Jocelia Grazia Co-Orientador: Prof. Dr. Cristiano F. Schwertner UNIVERSIDADE FEDERAL DO RIO GRANDE DO SUL PORTO ALEGRE 2014 “Sistemática de Thyreocoridae Amyot & Serville (Hemiptera: Heteroptera: Pentatomoidea): revisão de Alkindus Distant, morfologia do ovo de duas espécies de Galgupha Amyot & Serville e análise cladística de Corimelaena White, com considerações sobre a filogenia de Thyreocoridae, e morfologia do ovo de 16 espécies de Pentatomidae como exemplo de uso de caracteres de imaturos em filogenias” VIVIANA CAUDURO MATESCO Tese apresentada como parte dos requisitos para obtenção de grau de Doutor em Biologia Animal, área de concentração Biologia Comparada. ________________________________________ Prof. Dr. Augusto Ferrari (UFRGS) ________________________________________ Dra. Caroline Greve (CNPq ex-bolsista PDJ) ________________________________________ Prof. Dr. Cláudio José Barros de Carvalho (UFPR) ________________________________________ Profa. Dra. Jocelia Grazia (Orientadora) Porto Alegre, 05 de fevereiro de 2014. AGRADECIMENTOS À minha orientadora, Profa. Dra. Jocelia Grazia, pelos ensinamentos e por todas as oportunidades que me deu durante os treze anos em que estive no Laboratório de Entomologia Sistemática. Ao meu co-orientador, Prof. Dr. Cristiano Feldens Schwertner, pela troca de idéias. Ao Programa de Pós-Graduação em Biologia Animal e ao Departamento de Zoologia, pelo suporte oferecido e especialmente à Ana Paula, à Cintia e à Marina, pela ajuda nos momentos necessários. À CAPES, pela concessão da bolsa. Ao Prof. Luiz Campos e ao Filipe Michels, pela inestimável colaboração em dois dos artigos dessa tese. Ao Filipe, pelas co-autorias verdadeiramente compartilhadas, por dividir angústias e não esmorecer frente às decepções. À Aline Barcellos, pela amizade, pelo auxílio nas etapas iniciais do projeto e pela companhia na visita aos museus argentinos. A todos os meus maravilhosos e fantásticos colegas do Laboratório de Entomologia Sistemática, pelo auxílio constante, pelo apoio nos momentos difíceis e pelo agradável ambiente de trabalho. Em especial, ao Felipe Simões, por estar sempre pronto a ajudar e a oferecer apoio e incentivo. Especialmente aos “colegas de copo e de cruz”, Caroline Greve, Augusto Ferrari, Luciana Weiler e Kim Barão, pela amizade, pela paciência e pela inestimável ajuda em absolutamente todos os momentos. Ao Kim, também pela intensa troca de idéias e pela colaboração nos trabalhos no último semestre do doutorado. Ao Augusto, também pela grande ajuda na análise e interpretação dos dados filogenéticos. À Caroline Greve e ao Fernando Rocha, por dividirem as grandes angústias e as pequenas alegrias e pelos jantares etílico-vegetarianos. À minha mãe, por sempre me ajudar a cuidar da bicharada e por todo o suporte em casa. Aos cães do campus do Vale, pelos rabos balançantes todos os dias. Ao Guido, Lola, Bianca, Luigi, Shiva, Mingau, Mimi, Fiona, Jack, Dior e Cinza (in memoriam), pela companhia constante e por me lembrarem todos os dias qual a razão de levantar pela manhã. Especialmente à Juraci, por me ensinar que motivação é tudo na vida. iii SUMÁRIO Agradecimentos............................................................................................................... iii Sumário............................................................................................................................ iv Organização da tese........................................................................................................ vii Resumo.......................................................................................................................... viii Abstract............................................................................................................................. x Introdução geral............................................................................................................. 1 1. Sistemática de Hemiptera e seus subgrupos................................................... 1 2. Sistemática de Thyreocoridae Amyot & Serville........................................... 2 2.1. Caracterização e histórico............................................................. 2 2.2. Estudos morfológicos.................................................................... 6 2.3. Posição filogenética....................................................................... 9 2.4. Distribuição e diversidade........................................................... 10 2.5. Hábitos alimentares e importância econômica............................ 11 2.6. Estágios imaturos........................................................................ 12 2.7. Uso de caracteres de ovos em filogenias..................................... 14 Objetivos......................................................................................................................... 16 Material e Métodos......................................................................................................... 17 Estudo dos ovos e dos adultos............................................................................... 17 Análises cladísticas............................................................................................... 18 Resumo dos resultados................................................................................................... 20 Capítulo 1.............................................................................................................. 20 Capítulo 2.............................................................................................................. 20 Capítulo 3.............................................................................................................. 21 Capítulo 4.............................................................................................................. 22 Referências..................................................................................................................... 23 Capítulo 1 – Revision of the genus Alkindus Distant (Hemiptera: Heteroptera: Thyreocoridae: Corimelaeninae)……………………………………………………… 59 Abstract……...………………………………………………………………….. 59 Introduction……………………………………………………………………... 59 iv Material and methods…………………………………………………………… 60 Results…………………………………………………………………………... 61 Alkindus Distant, 1889…………….…………………………………………... 61 Key to the species of Alkindus….……………………………………………... 68 Alkindus atratus Distant, 1889………………………………………………… 68 Alkindus crassicosta Horvath, 1919…………………………………………... 69 References………………………………………………………………………. 70 Capítulo 2 – Egg ultrastructure of two species of Galgupha Amyot & Serville, with a discussion of the eggs and oviposition patterns of thyreocorid and allied groups (Hemiptera: Heteroptera: Pentatomoidea: Thyreocoridae)............................................ 73 Abstract…………………………………………………………………………. 73 Introduction........................................................................................................... 73 Material and methods............................................................................................ 74 Results and Discussion.......................................................................................... 74 Galgupha (Euryscytus) difficilis......................................................................... 74 Galgupha (Gyrocnemis) fossata......................................................................... 74 General morphology........................................................................................... 76 Oviposition behavior and maternal care............................................................. 77 Considerations on the relationships within Pentatomoidea…………………… 78 References............................................................................................................. 79 Capítulo 3 – Cladistic analysis of Corimelaena White based on morphology, with considerations on the phylogeny of the Thyreocoridae (Hemiptera: Heteroptera: Pentatomoidea)………………………………………………………………………... 82 Abstract……………………………………………………………………......... 82 Introduction……………………………………………………………………... 83 Material and Methods…………………………………………………………... 86 Taxon sampling……………………………………………………................... 86 Character sampling……………………………………………………………. 87 Cladistic analysis…………………………………………...…………………. 87 Results……………………………………………………………………........... 88 Cladistic analysis................................................................................................ 88 Characters……………………………………………………………… 88 v Trees………………………………………………………………….. 112 Discussion…………………............................................................................... 113 Conclusions......................................................................................................... 118 References..........................................................................................................
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