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MICROMORFOLOGIA E ANATOMIA FLORAL DAS SEÇÕES NEOTROPICAIS DE Bulbophyllum THOUARS (ORCHIDACEAE, ASPARAGALES): CONSIDERAÇÕES TAXONÔMICAS E EVOLUTIVAS

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MICROMORFOLOGIA E ANATOMIA FLORAL DAS SEÇÕES NEOTROPICAIS DE Bulbophyllum THOUARS (ORCHIDACEAE, ASPARAGALES): CONSIDERAÇÕES TAXONÔMICAS E EVOLUTIVAS UNIVERSIDADE ESTADUAL PAULISTA unesp “JÚLIO DE MESQUITA FILHO” INSTITUTO DE BIOCIÊNCIAS – RIO CLARO PROGRAMA DE PÓS-GRADUAÇÃO EM CIÊNCIAS BIOLÓGICAS (BIOLOGIA VEGETAL) MICROMORFOLOGIA E ANATOMIA FLORAL DAS SEÇÕES NEOTROPICAIS DE Bulbophyllum THOUARS (ORCHIDACEAE, ASPARAGALES): CONSIDERAÇÕES TAXONÔMICAS E EVOLUTIVAS ELAINE LOPES PEREIRA NUNES Tese apresentada ao Instituto de Biociências do Câmpus de Rio Claro, Universidade Estadual Paulista, como parte dos requisitos para obtenção do título de Doutor em Ciências Biológicas (Biologia Vegetal). Setembro - 2014 UNIVERSIDADE ESTADUAL PAULISTA unesp “JÚLIO DE MESQUITA FILHO” INSTITUTO DE BIOCIÊNCIAS – RIO CLARO PROGRAMA DE PÓS-GRADUAÇÃO EM CIÊNCIAS BIOLÓGICAS (BIOLOGIA VEGETAL) MICROMORFOLOGIA E ANATOMIA FLORAL DAS SEÇÕES NEOTROPICAIS DE Bulbophyllum THOUARS (ORCHIDACEAE, ASPARAGALES): CONSIDERAÇÕES TAXONÔMICAS E EVOLUTIVAS ELAINE LOPES PEREIRA NUNES Orientadora: Profa. Dra. Alessandra Ike Coan Coorientador: Prof. Dr. Eric de Camargo Smidt Setembro - 2014 581.4 Nunes, Elaine Lopes Pereira N972m Micromorfologia e anatomia floral das seções neotropicais de Bulbophyllum Thouars (Orchidaceae, Asparagales) : considerações taxonômicas e evolutivas / Elaine Lopes Pereira Nunes. - Rio Claro, 2014 252 f. : il., figs., tabs. Tese (doutorado) - Universidade Estadual Paulista, Instituto de Biociências de Rio Claro Orientador: Alessandra Ike Coan Coorientador: Eric de Camargo Smidt 1. Anatomia vegetal. 1. Anatomia floral de Orchidaceae. 3. Dendrobieae. 4. Epidendroideae. 5. Labelo. 6. Nectário. 6. Osmóforos. I. Título. Ficha Catalográfica elaborada pela STATI - Biblioteca da UNESP Campus de Rio Claro/SP Agradecimentos A Universidade Estadual Paulista “Júlio de Mesquita Filho” (UNESP, campus de Rio Claro), seu corpo docente e técnico-administrativo, por me receber de braços abertos e permitir a realização de um sonho. A Universidade Federal do Paraná (onde me graduei, me tornei mestre e servidora), seu corpo docente e técnico-administrativo, pela formação inicial e licença de três anos que permitiu que eu concluísse o tão almejado doutorado. A Ruhr-Universität Bochum, seu corpo docente, técnico-administrativo e discente, pela infraestrutura disponibilizada, todo o auxílio e prestatividade. À FAPESP pelas bolsas concedidas (de doutorado regular e BEPE - bolsa estágio em pesquisa no exterior), sem as quais esse trabalho não poderia ter sido desenvolvido. A Dra. Alessandra Ike Coan, pela confiança, pelos ensinamentos, por toda a paciência e pelas broncas mais que merecidas. Mas, sou grata principalmente pela orientação de forma tão presente, por vezes assumindo papel de psicóloga e amiga, mas sem nunca suprimir minhas ideias e tirar a identidade do trabalho. Ao Dr. Eric C. Smidt, pela coorientação, confiança, discussões e por me apresentar aos “Bulbophyllums”. Ao Dr. Thomas Stützel, por me receber no “Lehrstuhl für Evolution und Biodiversität der Pflanzen (Ruhr-Universität Bochum)”, por sua supervisão tão dedicada e aconselhamento. Ao grupo de colegas de pós e amigos carinhosamente apelidado de “in flames” (Kaire Nardi, Luís Felipe Daibes, Mariana Maciel, Naiara Lopes, Paula Maldonado e Thales Henrique Dias) por serem os melhores amigos “do Brasil e do Paraná”, pelo companheirismo, risadas e por serem minha família em Rio Claro. Aos colegas de laboratório (Ângela, Arthur, Blanca, Kaire, Letícia, Mariana, Mayra, Paula, Renata, Rita, Shirley, Thales, Yuri) pela convivência amigável e divertida, além da troca de experiências e aprendizados. A Paula Maldonado, pela parceria incrível no capítulo de B. sect. Micranthae, além de toda a ajuda com pranchas, formatação (além de me ajudar a manter a sanidade) nos relatórios e fim da tese. Ao Luís Felipe, meu vizinho, melhor amigo e irmão do coração por tudo (TUDO MESMO! A lista é muito grande e por vezes impublicável). Amo-te! Aos demais amigos e colegas da pós, companheiros de “salinha”, sujinhos e afins (Elizabeth - Beth, Bruna, Diogo, Eliana, Maria Gabriela - Gabi, Milene, Natália Costa, Nathália Miranda, Talita, Vítor), o meu muito obrigada por todos os momentos de alegria, devaneios, descontração, companheirismo, carinho e por aguentarem essa curitibana metida. A Missiani Ciochetta de Mello, colega na Ruhr que, além de companheira de laboratório, viagens e festas nos meus quatro meses de Alemanha, se tornou uma grande amiga. A Eliana e Milene, em especial pelo apoio durante o período na Alemanha. A amizade e presença de vocês (ainda que separadas por horas de trem) fizeram a estadia bem mais agradável. A minha família por ter me dado as ferramentas para realizar meus sonhos, por ter me apoiado durante a vida toda e compreendido as minhas muitas ausências. A todas as pessoas legais que conheci na cidade de Rio Claro e que fizeram com que eu deixasse de detestá-la e passasse e amá-la. Ao “quarteto fantástico” (Jenifer, Jocieli e Makeli), por estarem sempre presentes independente da distância. Um obrigada especial à Makeli, por ter me levado pra Botânica na graduação e, por ter me acolhido com tanto carinho em Piracicaba na transição entre Curitiba e Rio Claro. Aos amigos de Curitiba, que compreenderam minha ausência e torceram pelo meu sucesso. Agradeço também a todos que passaram pela minha vida ao longo destes quatro anos, enriquecendo-a e me ensinando algo. “Aqueles que passam por nós não vão sós, não nos deixam sós. Deixam um pouco de si, levam um pouco de nós”. Ao Lucas - last but not least – por te se juntado a minha caminhada num momento tão importante, por toda a compreensão enquanto eu “sofria de tese”, mas principalmente por todos os sorrisos e pelo amor. [...] Nobody said it was easy It's such a shame for us to part Nobody said it was easy No one ever said it would be this hard Oh, take me back to the start I was just guessing at numbers and figures Pulling the puzzles apart Questions of science, science and progress Do not speak as loud as my heart [...] (The Scientist – Coldplay) RESUMO Bulbophyllum, o maior gênero de Orchidaceae, possui distribuição pantropical mas sua maior diversidade concentra-se no sudeste Asiático. Na região Neotropical, o gênero é representado por 60 espécies, sendo que 49 delas ocorrem no Brasil. A maior diversidade de espécies neotropicais encontra-se no Sudeste brasileiro, nas regiões de contato entre o Cerrado e a Floresta Atlântica. Após o advento de filogenias moleculares, auxiliadas por dados morfológicos, foram reconhecidas e recircunscritas seis seções monofiléticas e neotropicais de Bulbophyllum: Bulbophyllaria, Furvescens, Napelli, Micranthae, Didactyle e Xiphizusa. Até o momento, informações sobre anatomia e micromorfologia florais para espécies neotropicais do gênero estão disponíveis apenas para seis espécies e tratam apenas do labelo. Além disso, informações sobre a polinização do grupo foram reportadas para apenas oito espécies. Considerando a recente recircunscrição do grupo, escassez de estudos sobre características florais e importância destas para a polinização em Orchidaceae, o objetivo deste trabalho foi analisar a micromorfologia e anatomia florais de 30 espécies neotropicais, bem como de sete espécies não-neotropicais e uma de Dendrobium, para: auxiliar na caracterização floral de cada seção, na diferenciação de espécies dentro das seções e para melhorar a compreensão do grupo como um todo. De forma geral, as flores das espécies das seções neotropicais de Bulbophyllum apresentam sépalas livres, com tricomas glandulares na face abaxial; pétalas com um feixe vascular colateral; labelo com epiderme estriada, com sulco na face adaxial, geralmente secretor, e uma quilha com estômatos na face abaxial; antera com epiderme papilosa e quatro polínias; presença de viscídio originado da desintegração das células do rostelo; ovário 6-lobado e com um feixe vascular por lobo. Cada seção apresenta características anatômicas e micromorfológicas florais comuns e outras que são variáveis e que são discutidas em cada capítulo, bem como suas implicações sistemáticas e para a polinização. Para diferenciar entre espécies de uma mesma seção, ornamentação de paredes periclinais externas e/ou cutícula das peças florais, número de feixes vasculares nas regiões medianas das peças florais e presença de tecidos secretores podem ser considerados características úteis. Na busca de possíveis sinapomorfias florais para o grupo Neotropical e para cada seção, apenas a presença de duas polínias em B. sect. Napelli e a presença de uma cavidade secretora no hipoquilo além de idioblastos com paredes espessadas no epiquilo em B. sect. Didactyle foram identificadas. Palavras-chave: Bulbophyllinae, Dendrobieae, Epidendroideae, labelo, nectário, osmóforo. ABSTRACT Bulbophyllum, the largest genus within Orchidaceae, is pantropical but its greatest diversity is found in the Asian Southeast. In the Neotropics, the genus comprises 60 species, being 49 of them found in Brazil. The greatest diversity of Neotropical species is in Brazilian Southeast, in transitional areas between the Atlantic Rainforest and Cerrado Vegetation. After the advent of molecular phylogenetic trees, and analysis of morphological data, six monophyletic Neotropical Bulbophyllum sections were recognised and recircumscribed: Bulbophyllaria, Furvescens, Napelli, Micranthae, Didactyle e Xiphizusa. However, information regarding floral anatomy and micromorphology for Neotropical species are available only for the labellum of six species. In addition, information on the pollination biology is available only to eight species. Considering the recent recircumscription of
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