Sistemática De Cyclanthaceae

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Sistemática De Cyclanthaceae Sistemática de Cyclanthaceae Eduardo da Silva Leal São Paulo 2018 Eduardo da Silva Leal Sistemática de Cyclanthaceae Systematics of Cyclanthaceae São Paulo 2018 Eduardo da Silva Leal Sistemática de Cyclanthaceae Systematics of Cyclanthaceae Tese apresentada ao Instituto de Biociências da Universidade de São Paulo, para obtenção do Título de Doutor em Ciências, na Área de Botânica. Orientador: Prof. Dr. Renato de Mello- Silva São Paulo 2018 da Silva Leal, Eduardo Sistemática de Cyclanthaceae. Orientador: Prof. Dr. Renato de Mello-Silva. 245 f. Tese (doutorado) – Instituto de Biociências da Universidade de São Paulo, Departamento de Botânica 1. Cyclanthaceae. 2. Dianthoveus. 3. Evodianthus. 4. Sistemática. 5. Taxonomia. Foto da capa: Juliana El Ottra 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 Dedico à Maitê Leal (Maitezoca), minha pequena filha, a qual espero que um dia supere à minha ausência nos últimos quatro anos. Agradecimentos Durante os últimos quatro anos muitas pessoas e instituições me proporcionaram condições necessárias para o desenvolvimento dessa tese. Sendo assim, expresso a minha mais profunda gratidão para cada um dos atores envolvidos nesse meu processo de formação. Ao meu orientador, Renato de Mello-Silva, por ter aceitado me orientar mesmo sem nunca ter me visto antes. O conhecimento compartilhado e a confiança depositada certamente me fizeram, e farão, ter um grande crescimento pessoal e profissional. À Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) pela concessão de bolsa de estudos durante os três primeiros anos de estudo; esse aporte financeiro foi fudamental para a execução do trabalho de campo e para a minha estadia em New York durante o doutorado sanduíche. À Universidade de São Paulo pela estrutura fornecida para o desenvolvimento da minha pesquisa e pelas oportunidades geradas para a minha constante qualificação. À Universidade Federal Rural da Amazônia pela concessão de licença para a realização do doutorado. À Rafaela Forzza, por sempre me motivar através de valorosas conversas, pelo incentivo e apoio durante esse trabalho; Ao Fabían Michelangeli por me receber tão amavelmente no New York Botanical Garden, e que, com paciência, acompanhou minhas primeiras incursões em um laboratório de Biologia Molecular. Aos professores do Laboratório de Sistemática Vegetal, profa. Lúcia Garcez Lohmann, prof. José Rubens Pirani e prof. Paulo Takeo Sano. Agradeço também aos funcionários do herbário Roberta Figueiredo, Viviane Jono e Abel Ribeiro pela presteza e auxílio. As pessoas que foram meus braços durante os primeiros dias de trabalho molecular, Lucas Bacci e Thuane Bochorny. A vocês serei eternamente grato. Em adição, também agradeço a Marcelo Reginato, Silvana Monteiro e Lucas Jordão pela companhia brasileira em solo estadunidense. Aos colegas e amigos de laboratório, de aprendizado, e por compartilhar memoráveis momentos de boas conversas, relaxamento e descontração: Andressa Cabral, Annelise Frazão, Beatriz Gomes, Benoît Loeiulle, Caetano Trancoso, Camila Monje, Carolina Siniscalchi, Caroline Andrino, Cíntia Luz, Daniela Costa, Eduardo Leal, Luana Sauthier, Luiz Fonseca, Eric Kataoka, Gisele Alves, Guilherme Antar, Jéssica Carvalho, Juan Pablo Nárvaez, Juliana El Ottra, Maila Beyer, Marcelo Devecchi, Marcelo Kubo, Mateus Cota, Matheus Santos, Matteus Colli, Maurício Watanabe, Paulo Gonella, Rebeca Viana, Rebeca Gama, Renato Ramos. Aos companheiros de campo Daniela Costa, Juliana El Ottra e em especial à Augusto Giaretta, companheiro em viagens para Colômbia e Guiana Francesa. Além disso, sua amizade, carinho e companheirismo o tornam ímpar e me fazem sentir orgulho de tê-lo como amigo. Agradeço também a Duban Canal, por me emprestar seu nome nas coletas colombianas, e Nils Köster por seu valoroso conhecimento e ótimo bom humor nas lindas e desafiadoras selvas da Colômbia. Ao outro Cyclanthólogo ativo, Dino Tuberquia. Obrigado pelo ensinamento e discussões sobre às Cyclanthaceae. Aos curadores dos herbários visitados, que me receberam com grande gentileza e atenção. À Mary Merello (MO) pelo envio de material da coleção viva do Missouri Botanical Garden. Agradeço ao meus pais, Moisés e Terezinha, minhas irmãs, Nircele e Roberta, cunhados Jean e Daniel e meus sobrinhos Heitor, Helena, Maysa, Arthur, Danilo, Thales e Laura, por todo carinho e amor dispensado ao longo da minha existência. À Tayná Machado por cuidar tão bem de nossa filha, Maitê Leal, que veio para dar um outro sentido à minha vida. Index General Introuction ............................................................................................... 03 Literature cited ...................................................................................................... 06 Chapter one: Phylogeny of Cyclantaceae (the Panama-hat family) based on molecular and morphological data ..................................................................... 09 Introduction ........................................................................................................... 10 Material & Methods .............................................................................................. 15 Results ................................................................................................................... 50 Discussion ............................................................................................................. 67 Conclusions ........................................................................................................... 76 Chapter two: Revisiting the Atlantic Forest cyclants with a key for the species and a revision of the endemic clade of Asplundia ............................................. 93 Introduction ........................................................................................................... 94 Material & Methods .............................................................................................. 95 Results ................................................................................................................... 96 Chapter three: Nomenclator botanicus of the Cyclanthaceae ....................... 125 Introduction ......................................................................................................... 126 Material & Methods ............................................................................................ 127 Results ................................................................................................................. 129 Final Considerations .......................................................................................... 237 Appendix ............................................................................................................. 239 Resumo Cyclanthaceae possui cerca de 230 espécies. São ervas terrestres, hemiepífitas, lianas ou epífitas distribuídas na região Neotropical, se estendendo do sul do México a Mata Atlântica do sul do Brasil. A família é dividida em duas subfamílias de acordo com a morfologia da inflorescência: Cyclanthoideae com flores estaminadas e pistiladas dispostas em ciclos alternos, incluindo apenas Cyclanthus; e Carludovicoideae com uma flor pistilada envolvida por quatro flores estaminadas e compreende o restante dos gêneros. Até o presente, estudos filogenéticos baseados somente em dados morfológicos, e com amostragem a nível genérico, não foram suficientes para clarificar suas relações. Além disso, um dos dois grandes grupos em Carludovicoideae emergiu como parafilético. Neste estudo, usamos dados morfológicos e de sete marcadores moleculares, cinco de cloroplastos e dois nucleares, com uma ampla amostragem à nível específico, para propor uma hipótese filogenética para Cyclanthaceae. Nosso resultado recuperou a família como monofilética e corroborou sua divisão em duas subfamílias. Os gêneros pertencentes a Carludovicoideae emergiram em três clados: (i) o gênero Carludovica; (ii) o clado Asplundia que contém Asplundia, Dicranopygium e Schultesiophytum; e (iii) o clado Evodianthus formado por Chorigyne, Dianthoveus, Evodianthus, Ludovia, Sphaeradenia, Stelestylis e Thoracocarpus. Todos os gêneros previamente circunscritos emergiram como monofiléticos. O grupo Sphaeradenia, composto por Chorigyne, Ludovia, Sphaeradenia e Stelestylis emerge como monofilético fortemente sustentado por caracteres morfológicos e moleculares. Para maximizar a informação filogenética, bem como facilitar a identificação, o gênero Dianthoveus é sinomizado em Evodianthus. No capítulo 2 uma chave de identificação para todas as espécies de Cyclanthaceae ocorrentes na Mata Atlântica é fornecida. As espécies endêmicas de Asplundia são revisadas, resultando em três táxons sinonimizados sob A. brachypus, e uma nova espécie, A. orthostigma, é descrita. No capítulo 3 é apresentado um índice taxonômico para todas as espécies já descritas de Cyclanthaceae. Nós indicamos 225 nomes aceitos e 60 sinônimos. Quatro lectótipos e dois neótipos são designados para quatro nomes. Para mais quatro nomes, a tipificação é completada, com a designação do segundo passo do lectótipo. Finalmente, doze nomes são considerados como novos sinônimos. Abstract. Cyclanthaceae has ca. 230 species. They are terrestrial herbs, hemiephytic, climbing or epiphytic distributed in the Neotropics, ranging from the South Mexico
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