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“Morfologia Comparada Da Flor De Melastomataceae Juss.” UNIVERSIDADE ESTADUAL DE CAMPINAS INSTITUTO DE BIOLOGIA JOÃO PAULO BASSO ALVES “MORFOLOGIA COMPARADA DA FLOR DE MELASTOMATACEAE JUSS.” CAMPINAS 2016 JOÃO PAULO BASSO ALVES “MORFOLOGIA COMPARADA DA FLOR DE MELASTOMATACEAE JUSS.” Tese apresentada ao Instituto de Biologia da Universidade Estadual de Campinas como parte dos reuisitos para a obtenção do Título de Doutor em Biologia Vegetal. ESTE ARQUIVO DIGITAL CORRESPONDE A VERSÃO FINAL DA TESE DEFENDIDA PELO ALUNO JOÃO PAULO BASSO ALVES E ORIENTADA PELA SIMONE DE PÁDUA TEIXEIRA. Orientadora: Prof. Dra. SIMONE DE PÁDUA TEIXEIRA Coorientador: Prof. Dr. RENATO GOLDENBERG CAMPINAS 2016 Campinas, 29 de junho de 2016. COMISSÃO EXAMINADORA Profa. Dra. Simone de Pádua Teixeira (Orientadora) Profa. Dra. Rosana Romero Profa. Dra. Priscila Andressa Cortez Profa. Dra. Maria das Graças Sajo Dra. Aline Oriani Os membros da Comissão Examinadora acima assinaram a Ata de Defesa, que se encontra no processo de vida acadêmica do aluno. Dedico a presente tese aos meus pais, Elza e João. “Vivendo, se aprende; mas o que se aprende, mais, é só a fazer maiores perguntas.” Guimarães Rosa; Grande Sertão: Veredas Agradecimentos À CAPES e a FAPESP (processo 2011/02701-0) pelo apoio financeiro a este trabalho. Ao Programa de Pós-Graduação em Biologia Vegetal do Instituto de Biologia da Unicamp. À Faculdade de Ciências Farmacêuticas de Ribeirão Preto por conceder a infraestrutura necessária à realização do trabalho. À Simone de Pádua Teixeira, pelo incentivo, inspiração e paciência. Muito Obrigado por esses dez anos de orientação e amizade! Ao Renato Goldenberg, pela disposição e confiança em coorientar o trabalho. À Ana Paula de Souza Caetano & Priscila Andressa Cortez, que tanto incentivaram e contribuíram com o projeto, do início ao fim. Muito Obrigado por me apresentarem às Melastomataceae! A todos que, de uma forma ou outra, contribuíram para as coletas realizadas durante meu doutorado: João Batista Alves, Ana Paula Caetano, Juliana Milani, Priscila Cortez, Renato Goldenberg, André Fontana, Marcelo Monge, Paola Ferreira, Eduardo Almeida, Juliana Paulino, Sidnei Mateus, Túlio Nunes, Lucas & Melina Torati, Fernanda Vergamini. À Adriane Cristina Sarti Sprogis e Stella Maris Fick de Ferraz (IB/Unicamp), Edimárcio da Silva Campos (FCFRP/USP), José Augusto Maulin (FMRP-USP) e Rodrigo Ferreira Silva (DQ-FFCLRP/USP) pelo apoio técnico e solicitude. À Maria Roseli de Melo, pela ajuda tranquilizadora com os trâmites institucionais da Unicamp. Ao Renato Goldenberg e ao Fabián Michelangeli, por compartilharem seu conhecimento e entusiasmo sobre Melastomataceae, além de sua vasta bibliografia. Aos amigos melastomatógos, participantes da disciplina oferecida em Campinas (2012/2015), pela colaboração constante. Ao Eduardo Almeida, pela generosidade em discutir os meandros da Biologia Comparada, em disciplinas e cafés. Ao Juca pelas sugestões sobre análise histológica. À Thais pela parceria durante o doutoramento. À Juliana Lischka Sampaio Mayer, Letícia Silva Souto e Rosana Romero, membros da banca de qualificação; e Aline Oriani, Ana Paula de Souza Caetano e Maria do Carmo Estanislau do Amaral, membros da pré-banca; pelas críticas e sugestões. Aos membros titulares e suplentes da banca examinadora, pela disponibilidade e disposição em contribuir com o trabalho. Aos amigos do Laboratório de Botânica, Aline Rejane, Ana Paula Caetano, Camila Devicaro, Cristina Marinho, Edimárcio Campos, Flávia Leme, Giseli Pedersoli, Isabel Nascimento, Juca San Martin, Juliana Milani, Juliana Paulino, Liana Capucho, Marina Costa, Priscila Cortez, Thais de-Barros e Viviane Leite, pelo auxílio e companheirismo ao longo desses anos de trabalho. Aos amigos da Unicamp, em especial André Scatigna, Gustavo Shimizu e Marcelo Monge pela prontidão e hospitalidade. Aos amigos, de perto e de longe, Fernanda, Gisele P, Lucas J, Melina, Marina, Pedro, Sarah, Sibila, Sidnei, Thiago, por estarem sempre presentes. Aos meus “irmãos”, Tati e Sandro, pelo apoio incondicional e por me ensinarem a ser no mundo. Aos meus pais, Elza e João Batista, por seu amor e amparo. Muito Obrigado por estarem ao meu lado. Faço um agradecimento especial à minha mãe, que tanto me ajudou ao final desse processo. Resumo Melastomataceae é o maior grupo de Myrtales, com cerca de 5000 espécies, distribuídas principalmente no Neotrópico. Apesar da família ser monofilética, seus membros exibem grande heterogeneidade morfológica, o que dificulta o estabelecimento de sinapomorfias, especialmente em termos de caracteres florais. O objetivo geral da presente tese foi comparar a morfologia da flor em desenvolvimento em algumas espécies de Melastomataceae, a fim de elucidar aspectos estruturais importantes para o grupo, como o merisma floral, posição do ovário e morfologia estaminal. Para tal, botões em diversos estádios de desenvolvimento de Henriettea saldanhae, Leandra melastomoides, Miconia dodecandra, Microlicia euphorbioides, Rhynchanthera grandiflora e Tibouchina grandiflora foram coletados, fixados em solução de Karnovsky ou FGAA e preparados para observações de superfície em microscopia eletrônica de varredura, histológicas em microscopia de luz e diafanizadas. Os resultados obtidos são apresentados em quatro capítulos que atendem objetivos específicos: (1) Compara a estrutura e vascularização floral das espécies mencionadas, acrescidas de Pleiochiton ebracteatum e Rupestrea johnwurdackiana; (2) Analisa as bases ontogenéticas para a variação da posição do ovário em Melastomataceae (que pode variar de súpero, semi-ínfero ou ínfero); (3) Estuda a diversidade estaminal em Melastomataceae a partir de um enfoque ontogenético; e (4) Elucida a estrutura do cálice em Leandra melastomoides, condição frequentemente chamada de “cálice duplo” por apresentar duas partes distintas e opostas. Os dados obtidos em cada capítulo são discutidos em um contexto filogenético, que podem ser extrapolados para outras espécies de Melastomataceae e Myrtales. Espera-se com isso contribuir com a sistematização de questões em Morfologia floral para novos estudos em Melastomataceae. Palavras-chave: anatomia vascular, hipanto, merisma, morfologia estaminal, Myrtales, ontogenia floral, posição do ovário, Vorläuferspitze. Abstract Melastomataceae is the largest group of Myrtales, with about 5000 species, mainly distributed in the Neotropics. Although the family is monophyletic, its members exhibit great morphological heterogeneity, which hinders the establishment of synapomorphies, especially in terms of floral characters. The overall objective of this thesis was to compare the flower anatomy and development in some species of Melastomataceae in order to elucidate important structural aspects for the group. For this, flowers and buds of several developmental stages of Henriettea saldanhae, Leandra melastomoides, Miconia dodecandra, Microlicia euphorbioides, Rhynchanthera grandiflora and Tibouchina grandiflora were collected, fixed in Karnovsky solution or FGAA and prepared for observations under light and scanning microscopy. The results are presented in four chapters that with objectives: (1) To compare the floral structure and vasculature of the species mentioned, plus Pleiochiton ebracteatum and Rupestrea johnwurdackiana; (2) To analyze the ontogenetic basis for the variation of the ovary position in Melastomataceae (in which the ovary varies from superior, semi-inferior or inferior); (3) To study the stamen morphology diversity in Melastomataceae from an ontogenetic approach; and (4) To elucidate the “double calyx” in Leandra melastomoides, which displays two distinct opposite parts. The data obtained for each chapter are discussed in a phylogenetic framework, which can be extrapolated to other Melastomataceae and Myrtales species. The results will contribute to the systematization of issues in floral morphology for further studies in Melastomataceae. Key words: vascular anatomy, hypanthium, stamen morphology, Myrtales, floral ontogeny, ovary position, Vorläuferspitze. Sumário Introdução Geral .................................................................................................................... 14 Sobre Morfologia Floral .................................................................................................... 20 Objetivos Gerais ................................................................................................................. 23 Estrutura da Tese ............................................................................................................... 23 Apêndice: Considerações sobre uma Terminologia para Melastomataceae ............ 25 Bibliografia:......................................................................................................................... 28 Capítulo 1. Estrutura floral comparada de Melastomataceae ......................................... 40 1.1. Resumo ..................................................................................................................... 40 1.2. Introdução ................................................................................................................ 41 1.3. Objetivos ................................................................................................................... 43 1.4. Materiais & métodos............................................................................................... 44 1.5. Resultados ................................................................................................................ 47 1.5.1. Henriettea saldanhae Cogn. (Henrietteeae)
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