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Is Pseudoxandra Spiritus-Sancti (Annonaceae), with Its Male and Bisexual Flowers, an Androdioecious Species?

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Is Pseudoxandra Spiritus-Sancti (Annonaceae), with Its Male and Bisexual Flowers, an Androdioecious Species? Filogenômica, morfologia e taxonomia na tribo Malmeeae (Malmeoideae, Annonaceae): implicações na evolução da androdioicia Phylogenomics, morphology and taxonomy in tribe Malmeeae (Malmeoideae, Annonaceae): implications on the evolution of androdioecy Jenifer de Carvalho Lopes São Paulo 2016 Filogenômica, morfologia e taxonomia na tribo Malmeeae (Malmeoideae, Annonaceae): implicações na evolução da androdioicia Phylogenomics, morphology and taxonomy in tribe Malmeeae (Malmeoideae, Annonaceae): implications on the evolution of androdioecy Jenifer de Carvalho Lopes Tese apresentada ao Instituto de Biociências da Universidade de São Paulo, para a obtenção de Título de Doutor em Ciências, na Área de Botânica. Orientador: Renato de Mello-Silva São Paulo 2016 Lopes, Jenifer de Carvalho Filogenômica, morfologia e taxonomia na tribo Malmeeae (Malmeoideae, Annonaceae): implicações na evolução da androdioicia / 98 Tese (Doutorado) – Instituto de Biociências da Universidade de São Paulo. Departamento de Botânica 1. Androdioicia. 2. Annonaceae. 3. Filogenômica. I. Universidade de São Paulo. Instituto de Biociências. Departamento de Botânica. Comissão Julgadora: ________________________ _______________________ Prof(a). Dr(a). Prof(a). Dr(a). ________________________ _______________________ Prof(a). Dr(a). Prof(a). Dr(a). ______________________ Prof. Dr. Renato de Mello-Silva Orientador Metafísica? Que metafísica têm aquelas árvores? A de serem verdes e copadas e de terem ramos E a de dar fruto na sua hora, o que não nos faz pensar, A nós, que não sabemos dar por elas. Mas que melhor metafísica que a delas, Que é a de não saber para que vivem Nem saber que o não sabem? Alberto Caeiro, heterônimo de Fernando Pessoa Trecho de Guardador de Rebanhos, poema quinto Agradecimentos Gostaria de agradecer à Universidade de São Paulo pela estrutura fornecida para o desenvolvimento da minha pesquisa e formação acadêmica. À Wageningen University e ao Royal Botanic Gardens, Kew, por me possibilitarem realizar minha pesquisa no exteiror. Agradeço também ao Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) pela bolsa de estudos concedida durante o primeiro ano do meu doutorado. À Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) pela bolsa de estudos nos demais anos do doutorado (2013/04902-9), pela Bolsa Estágio Pesquisa no Exterior (2014/09366-0) que financiou minha ida e manutenção na Holanda e Inglaterra para desenvolvimento de minha pesquisa e pelo auxílio regular à pesquisa concedido ao meu orientador, professor Renato de Mello-Silva, (2014/09131-3), que custeou as despesas com minha pesquisa tanto na Holanda quanto na Inglaterra. A Alberta Mennega Foudation, Holanda, que me contemplou com um auxílio financeiro que custeou parte das despesas laboratoriais em Wageningen. Meus mais sinceros agradecimentos ao professor Renato de Mello-Silva por esses nove anos que trabalhamos juntos, contando iniciação científica, mestrado e doutorado. Agradeço à paciência com que me ensinou desde a escrever melhor, incluindo o uso da crase, a ter acurácia no desenvolvimento da minha pesquisa, a ter cuidado e zelo pela coleção do herbário e às inumeráveis outras coisas que aprendi com sua exemplar conduta profissional. Agradeço também aos professores do laboratório de Sistemática Vegetal, profa. Lúcia Garcez Lohmann, prof. José Rubens Pirani e prof. Paulo Sano, aos funcionários Abel Ribeiro, Roberta Figueiredo e Viviane Jono, aos meus colegas Alexandre Zuntini, Augusto Giaretta, Beatriz Machado, Carolina Andrino, Carolina Sinischalchi, Cíntia Luz, Daniela Costa, Eduardo Leal, Eric Kataoka, Euder Martins, Guilherme Antar, Juan Pablo Narvaéz, Juliana El’Ottra, Leonardo Borges, Marcelo Devecchi, Marcelo Kubo, Matheus Martins, Mayla Beyer, Paulo Gonella, Rebeca Viana, Renato Ramos, Verônica Thode pelo apoio, em especial a Gisele Alves e Jéssica Nayara Francisco; a Dra. Juliana Lovo pelo auxílio no desenvolvimento do meu projeto de anatomia floral; a Annelise Frazão pela ajuda nas análises filogenéticas e a Luiz Henrique Fonseca pelo auxílio nas análises de evolução de estados ancestrais. Agradeço a Dr. Lars Chatrou, por me receber na Wageningen University, por me passar seu vasto conhecimento sobre sistemática e pela cordialidade durante o um ano que estive na Holanda. Agradeço também a Dra. Sabrina Simon pela tradução do alemão para o inglês dos comentários dos protólogos das espécies de Ephedranthus. A todos os meus colegas do Biosystematic Group, em especial a Lidja Berke, Sara van der Kerke, Zhen Wei e Paul Hoekstra. Agradeço a Dr. Paul Maas e a senhora Hiltje Maas por me darem carona inúmeras vezes até o herbário em Leiden e pela hospitalidade. Agradeço a Roy Erkens por me convidar a participar do workshop sobre filogenômica de Annonaceae realizado na China, durante o período que estive na Holanda, ao qual tive as despesas de transporte e de hospedagem pagas por seu projeto. Agradeço também a Dra. Paula Rudall por me receber no Jodrell Laboratory no Royal Botanic Gardens, Kew e a Dra. Maria Graça Sajo pelo auxílio na execução do meu projeto com anatomia floral. Agradeço aos meus familiares, minha mãe, minha tia Elena, meu irmão Denis e meu marido, Vítor Dantas, pelo apoio incondicional. Índice Resumo. ....................................................................................................................................... 1 Abstract. ...................................................................................................................................... 2 Introdução ................................................................................................................................... 3 Referências bibliográficas ...................................................................................................... 6 Capítulo 1 .................................................................................................................................... 9 Phylogenomics and evolution of androdioecy in tribe Malmeeae (Annonaceae), a Neotropical woody group ........................................................................................................... 9 Abstract. .................................................................................................................................. 9 Introduction .......................................................................................................................... 10 Material & Methods ............................................................................................................. 11 Results .................................................................................................................................... 23 Discussion .............................................................................................................................. 32 Conclusions ........................................................................................................................... 41 References.............................................................................................................................. 42 Capítulo 2 .................................................................................................................................. 49 Is Pseudoxandra spiritus-sancti (Annonaceae), with its male and bisexual flowers, an androdioecious species? ...................................................................................................... 49 Abstract ................................................................................................................................. 49 Introduction .......................................................................................................................... 50 Material & Methods ............................................................................................................. 51 Results .................................................................................................................................... 54 Discussion .............................................................................................................................. 56 Conclusions ........................................................................................................................... 60 References.............................................................................................................................. 61 Capítulo 3 .................................................................................................................................. 64 Morphological phylogenetic analysis and revision of Ephedranthus (Annonaceae) .......... 64 Abstract ................................................................................................................................. 64 Introduction .......................................................................................................................... 65 Material & Methods ............................................................................................................. 66 Results .................................................................................................................................... 70 Phylogenetic relationships in Ephedranthus ...................................................................... 70 Taxonomic Treatment .......................................................................................................... 72 Key to the species .................................................................................................................
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