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Taxonomy, Phylogeny and Evolution of Egg- Powdering Behavior Associated Traits in Molomea China, 1827 (Insecta: Hemiptera: Cicadellidae)

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Taxonomy, Phylogeny and Evolution of Egg- Powdering Behavior Associated Traits in Molomea China, 1827 (Insecta: Hemiptera: Cicadellidae) TAXONOMY, PHYLOGENY AND EVOLUTION OF EGG- POWDERING BEHAVIOR ASSOCIATED TRAITS IN MOLOMEA CHINA, 1827 (INSECTA: HEMIPTERA: CICADELLIDAE) Beatriz Macharet Camisão de Vasconcelos Dissertação de Mestrado apresentada ao Programa de Pós-graduação em Biodiversidade e Biologia Evolutiva, Instituto de Biologia, Universidade Federal do Rio de Janeiro, como parte dos requisitos necessários à obtenção do título de Mestre em Biodiversidade e Biologia Evolutiva. Orientadora: Profa. Dra. Daniela M. Takiya Rio de Janeiro Janeiro de 2014 i TAXONOMY, PHYLOGENY, AND EVOLUTION OF EGG-POWDERING BEHAVIOR ASSOCIATED TRAITS IN MOLOMEA CHINA, 1827 (INSECTA: HEMIPTERA: CICADELLIDAE) Beatriz Macharet Camisão de Vasconcelos Dissertação de Mestrado submetida ao Programa de Pós-graduação em Biodiversidade e Biologia Evolutiva, Instituto de Biologia, Universidade Federal do Rio de Janeiro, como parte dos requisitos necessários à obtenção do título de Mestre em Biodiversidade e Biologia Evolutiva. Orientadora: Profa. Dra. Daniela Maeda Takiya Rio de Janeiro Janeiro de 2014 ii TAXONOMY, PHYLOGENY, AND EVOLUTION OF EGG-POWDERING BEHAVIOR ASSOCIATED TRAITS IN MOLOMEA CHINA, 1827 (INSECTA: HEMIPTERA: CICADELLIDAE) Beatriz Macharet Camisão de Vasconcelos Orientadora: Profa. Dra. Daniela Maeda Takiya Dissertação de Mestrado submetida ao Programa de Pós-graduação em Biodiversidade e Biologia Evolutiva, Instituto de Biologia, Universidade Federal do Rio de Janeiro, como parte dos requisitos necessários à obtenção do título de Mestre em Biodiversidade e Biologia Evolutiva. Aprovada por: ___________________________________________________________ Presidente: Prof. Dr. José Ricardo M. Mermudes (Instituto de Biologia, UFRJ) ___________________________________________ Prof. Dr. Gabriel Mejdalani (Museu Nacional, UFRJ) _______________________________ Prof. Dr. Christopher H. Dietrich (University of Illinois at Urbana Champaign, EUA) _______________________________ Suplente: Profa. Dra. Cássia M. Sakuragi (Instituto de Biologia, UFRJ) _______________________________ Suplente: Dra. Olívia Evangelista (Museu de Zoologia, USP) Rio de Janeiro Janeiro de 2014 iii FICHA CATALOGRÁFICA VASCONCELOS, Beatriz Macharet Camisão de. Taxonomy, phylogeny, and evolution of egg-powdering behavior associated traits in Molomea China, 1827 (Insecta: Hemiptera: Cicadellidae) – Beatriz Macharet Camisão de Vasconcelos - Rio de Janeiro: UFRJ/ Instituto de Biologia, 2014. (xx-xx-XX f. il.). Orientador: Profa. Dra. Daniela Maeda Takiya Dissertação (mestrado) – UFRJ/ Instituto de Biologia/ Programa de Pós-graduação em Biodiversidade e Biologia Evolutiva, 2014. Referências Bibliográficas: f. 100-103. 1. Insecta. 2. Hemiptera. 3. Cicadellidae. 4. Molomea China, 1927. 5. Taxonomia. 6. Filogenia. I. Daniela Maeda Takiya II.Universidade Federal do Rio de Janeiro, Intituto de Biologia, Programa de Pós-graduação em Biodiversidade e Biologia Evolutiva. III. Taxonomy and Phylogeny of the genus Molomea China, 1927 (Insecta: Hemiptera: Cicadellidae). iv To my dear aunt Celi, a great enthusiast, supporter and sponsorer of the study of leafhoopers. v ACKNOWLEDGEMENTS First of all, I must thank my supervisor, Prof. Dr. Daniela M. Takiya, for the great patience, the attention to details, dedication, and for teaching me so much since I first started studying leafhoppers. She never gave up on me, even though I drove her mad sometimes. I just wish I could be half the teacher that you are. To the examiners, for accepting to participate and contribute to my work. To Prof. Jorge Nessimian, Prof. Nelson Ferreira Jr. and Prof. José Ricardo Mermudes, and all the other members of the Laboratório de Entomologia, for becoming almost my second family and for filling my days with laughter, but also helping me with my innumerous questions and doubts when the occasion was serious. Special thanks to Clayton Gonçalves for helping me with photographs and giving me tips about my presentations and my plates, and of course for all the funny jokes. To my family, who never questioned my choices, even though they don’t understand them completely sometimes. Thank you for all the love and support. I must thank my friends from school, Paula Belchior, Raisa Belchior, Raizza Teixeira, Rodrigo Hull, Susana Resende, Tatiana Póvoa, Loamy Cavalcanti and Bruna Marques. You are my oldest friends, and you know me better than anyone else. We’ve been through a lot of changes, but I know that no matter what happens with us, we will always love each other. Thank you for being in my life for so long. To my goose friends, Giulia Engel, Juliana Valle, Luana Calazans and Rafaella Garbin, for all the long talks on the cellphone, all the jokes and for hearing me complain about life so many times. I miss our classes together. I must thank also my good friends Ana Luiza Pimenta, Manuella Folly and Tainá Stauffer. Thank you for all the drinks, the talks, the advices, the trips, and for letting me know that I can always count on you. Special thanks to Pedro Fasura, who has experienced very closely every moment of this work, both good and bad. Thank you for all the attention, the partnership, the help, and for filling my days with joy, kindness and affection. You make me incredibly happy. Finally I must thank Dr. Jürgen Deckert and Dr. Roland Mühlethaler for helping me during my visit to the Museum für Naturkunde, Berlin, and also Dr. Wolfgang vi Nässig, Dr. Schröder and his wife for helping me during my visit to the Senckenberg Museum in Frankfurt, Germany. Financial support was given by CNPq/PROTAX. vii “Nothing in Biology makes sense except in the light of Evolution" Theodosius Dobzhansky, 1973 “What makes things baffling is their degree of complexity, not their sheer size…a star is simpler than an insect” Martin Rees, 1999 “Science never pursues the illusory aim of making its answers final, or even probable. Its advance is, rather, towards the infinite yet attainable aim of ever discovering new, deeper, and more general problems, and of subjecting its ever tentative answers to ever renewed and ever more rigorous tests. The wrong view of science betrays itself in the craving to be right; for it is not his possession of knowledge, of irrefutable truth, that makes the man of science, but his persistent and recklessly critical quest for truth” Karl Popper, 1959 viii ABSTRACT TAXONOMY, PHYLOGENY AND EVOLUTION OF EGG-POWDERING BEHAVIOR ASSOCIATED TRAITS IN MOLOMEA CHINA, 1827 (INSECTA: HEMIPTERA: CICADELLIDAE) Beatriz Macharet Camisão de Vasconcelos Advisor: Dr. Daniela M. Takiya The twenty three known species of Molomea China are recorded from Argentina, Brazil, Ecuador, Paraguay and Peru. The identification of the species and the only taxonomic key to the genus are based on characters of the male, and the female terminalia was never studied in detail. Females of Molomea cover the incisions made in the plant tissue for oviposition with brochosomes, and this behavior is associated with two dimorphic features that are not always found in species among the genus. The aim of this study was to propose and discuss phylogenetic hypothesis for the species of Molomea based on morphological and molecular data, to describe the female terminalia, to provide a taxonomic key for males and females and also to discuss the patterns of gains and losses of the dimorphic characters related to the egg-powdering behavior in Molomea. A new species of Molomea from Rio de Janeiro, Brazil, is described and illustrated. Two females of Molomea that probably belong to two new species of the genus are described and illustrated. The type specimens of M. alternata, and M. laminata were described and illustrated, as the syntypes of M. vermiculata, which was found to belong to a different genus. The males of M. guttuata and M. infulata, known only from female types, are described and illustrated. A polymorphism of the male genitalia was illustrated and discussed. A comparative study of the female terminalia is showed. Eighty-three morphological characters were coded and the parsimony and Bayesian analyses found the genus monophyletic. The molecular analyses of four genes (16S, COI, COII and H3) individually and combined with maximum likelihood, and Bayesian inference obtained similar results, however the parsimony analyses of the same datasets obtained different topologies. Neither of the tree approaches supported ix the monophyletism of Molomea. The morphological and molecular datasets were also analyzed together with parsimony, maximum likelihood and Bayesian inference, what helped to solve the relationships of some internal branches. The efficiency of the molecular data is discussed. The two dimorphic characters associated with the egg- powdering behavior were optimized on the most parsimonious trees, and it was found that the region with distinctly concentrate setae on female forewing originated at least seven times inside the genus, while the elongate setae on female hindtibia was a characteristic already found in the ancestral of the genus that was lost independently three times inside Molomea. A S-DIVA analysis was performed and it was found that the losses and gains of these two dimorphic characters in Molomea cannot be related to shifts to major phytogeographical regions. x RESUMO TAXONOMIA, FILOGENIA E EVOLUÇÃO DOS CARACTERES ASSOCIADOS AO COMPORTAMENTO DE OVIPOSIÇÃO EM MOLOMEA CHINA, 1927 (INSECTA: HEMIPTERA: CICADELLIDAE) Beatriz Macharet Camisão de Vasconcelos Orientadora: Dr. Daniela M. Takiya As 23 espécies de Molomea China, 1927 estão distribuídas na Argentina, Brasil, Equador,
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