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Sistemática De Drosera Sect. Drosera Ss (Droseraceae)

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Sistemática De Drosera Sect. Drosera Ss (Droseraceae) Paulo Sergio Minatel Gonella Silva Sistemática de Drosera sect. Drosera s.s. (Droseraceae) Systematics of Drosera sect. Drosera s.s. (Droseraceae) São Paulo 2017 ! ! Sistemática de Drosera sect. Drosera s.s. (Droseraceae) Systematics of Drosera sect. Drosera s.s. (Droseraceae) Paulo Minatel Gonella Tese apresentada ao Instituto de Biociências da Universidade de São Paulo para a obtenção de Título de Doutor em Ciências Biológicas na Área de Botânica. Orientador Paulo Takeo Sano (Universidade de São Paulo) Co-orientador Andreas Fleischmann (Botanische Staatssammlung München) São Paulo — 2017 ! Ficha Catalográfica Gonella, Paulo Minatel Sistemática de Drosera sect. Drosera s.s. (Droseraceae) 159 pg. Tese (Doutorado) – Instituto de Biociências da Universidade de São Paulo. Departamento de Botânica. 1. Droseraceae. 2. Sistemática. 3. Diversidade I. Universidade de São Paulo. Instituto de Biociências. Departamento de Botânica. Comissão Julgadora Prof. Dr. Paulo Takeo Sano (Orientador) : Prof(a). Dr(a). : Prof(a). Dr(a). : Prof(a). Dr(a). : Prof(a). Dr(a). : ! Nas muitas curvas sinuosas que a vida dá é certo que encontremos obstáculos, alegrias, tristezas, quiçá até mesmo uma espécie nova no Facebook, mas nenhum desses achados é mais valioso que uma amizade verdadeira. Aos meus queridos Carol e Baladinha, dedico. ! ! Só a Natureza é divina, e ela não é divina... Se às vezes falo dela como de um ente É que para falar dela preciso usar da linguagem dos homens Que dá personalidade às cousas, E impõe nome às cousas. Mas as cousas não têm nome nem personalidade: Existem, e o céu é grande e a terra larga, E o nosso coração do tamanho de um punho fechado... Bendito seja eu por tudo quanto não sei. É isso tudo que verdadeiramente sou. Gozo tudo isso como quem sabe que há o sol. Alberto Caeiro O Guardador de Rebanhos ! ! Agradecimentos A construção de um doutor é feita de muito trabalho, dedicação, força de vontade e vários outros sacrifícios individuais. No entanto, o mérito por esse trabalho e pela minha formação pessoal e acadêmica não é só meu, mas de todos que me ajudaram a trilhar esse caminho. Nesses quatro anos, tive apoio e suporte de diversas pessoas e instituições e aproveito esse espaço para agradecê-las. Agradeço às instituições que ampararam esse doutorado: Instituto de Biociências da Universidade de São Paulo, Ludwig Maximilians Universität München, Botanische Staatssammlung München, Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq – Proc. 140135/2013-8) e Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES – PDSE/Proc. 99999.010612/2014-09). Vou quebrar o protocolo e agradecer em primeiro lugar aos meus orientadores desde sempre, meus pais Nadir e Sérgio, que sempre estiveram lá para me guiar, apoiar e dar aquele empurrãozinho quando necessário. Obrigado por tudo! Ao meu orientador, Paulo Sano, que me acompanha desde a graduação, agradeço por todo apoio, aprendizado e orientação não só acadêmica, mas também pessoal ao longo desses anos todos. Ao meu co-orientador, Andreas Fleischmann, pela recepção calorosa em Munique e pelas incontáveis horas de aprendizado, seja no laboratório, no Jardim Botânico, em campo ou num Biergarten. Aos que contribuíram com o desenvolvimento da minha pesquisa, em especial ao Fernando Rivadavia pelas incontáveis horas online e no campo de frutíferas discussões sobre as dróseras. Ao Dr. David Neill pela sua contribuição no Capítulo 2. Ao Carlos Rohrbacher, que contribuiu com material e importantes observações sobre as plantas em seu excepcional cultivo. Ao Reginaldo Vasconcelos, o descobridor da Drosera magnifica, por toda sua contribuição para que pudéssemos estudar essa magnífica planta e pelos seus esforços em protege- la. Agradeço aos meus amigos e companheiros, atuais e antigos, do Sobre-as- Ondas, Annelise Frazão, Augusto Giaretta, Beatriz Gomes, Caetano Troncoso, Carolina Siniscalchi, Carolina Agostini, Caroline Andrino, Cíntia Luz, Eric ! Kataoka, Fernanda Calió, Gisele Alves, Guilherme Antar, Gustavo Heiden, Jenifer Carvalho, Juliana Rando, Juliana El Otra, Juliana Lovo, Leonardo Borges, Livia Echternacht, Luiz Henrique Fonseca, Marcelo Devecchi, Marcelo Kubo, Matheus Cota, Pamela Santana, Paulo Baleeiro, Pedro Fiaschi e Rebeca Viana. Um agradecimento especial à Anne, à Beca, ao Caqui, à Carol e ao Kubo que foram maravilhosos sempre, mas em especial por segurarem a minha barra na reta final dessa fase. Aos colegas e amigos do AG Heubl em Munique, Prof. Günther Heubl, Agnes Scheunert, Andrea Brandl, Tanja Ernst, Salvatore Tomasello e, claro, Andreas Fleischmann, pela companhia e aprendizado. Um agradecimento especial ao Prof. Heubl, que financiou os custos laboratoriais envolvidos na obtenção dos dados do Capítulo 4. Aos professores que fizeram parte da minha banca de qualificação, Prof. José Rubens Pirani e Profa. Fernanda Calió, pelas valiosas sugestões. Aos técnicos do Herbário SPF, Viviane Jono, Roberta Figueiredo e Abel Cangussu pelo zelo com essa preciosa coleção e pelo café de todo dia. Aos amigos que me receberam na Alemanha, em especial ao Alexander Schommer, Juilson Jubanski, Jan Zippel, Denison Caldeiron, Tiago de Oliveira, Roberto Goya, Alexander Rockinger e Thomas Wittmann, que tornaram meu tempo na batatolândia ainda mais memorável. Aos meus amigos dos tempos de graduação, Caroline “Cidoca” Araujo, Ivan Dias, Victor Rabelo, Diego Vasques e Yve Magedans, pelo incentivo e companheirismo. À minha tia Vanda, pela companhia sempre especial. À minha tia Teresinha, que sempre me incentivou. À minha avó Sônia, pela sua alegria contagiante. À minha avó Marina, que me ensinou a ver a beleza das flores e ao meu avô Guilherme, que me levou pra vê-las na natureza. E, mais uma vez porém nunca o suficiente, a essas pessoas lindas que são a Carol, a Sinis, o Baladinha e o Kubo, com quem dividi momentos maravilhosos durante esse doutorado e que tornaram esse processo de crescimento acadêmico, mas, sobretudo, pessoal, muito mais fácil e especial. Muito obrigado! Thank you very much! Danke schön! Paulo ! Contents Abstract ..................................................................................................................... 1 Resumo ..................................................................................................................... 2 General Introduction ................................................................................................ 3 Chapter 1 — Drosera magnifica: the largest New World sundew, discovered on Facebook .................................................................................................................... 9 Chapter 2 — A revision of Drosera (Droseraceae) from the central and northern Andes, including a new species from the Cordillera del Cóndor (Peru and Ecuador) ................................................................................................................................. 23 Chapter 3 — A synopsis of the genus Drosera (Droseraceae) from Brazil ............ 39 Chapter 4 — Phylogenetic Analyses of Neotropical Drosera (Droseraceae) ....... 127 General Conclusions .......................................................................................... 157 List of author’s publications .............................................................................. 159 ! ! Abstract Drosera is a carnivorous plant genus belonging to the Droseraceae and comprising around 250 species distributed worldwide. A great number of new species and combinations have been published for Brazil and the Neotropics in the past decades; however, a comprehensive taxonomic work dealing with these species was still lacking. Furthermore, previous phylogenetic works on the genus focused on understanding the relationships between the morphological and geographical groups, revealing that all proposed infrageneric classifications were incongruent with the phylogeny and that the Neotropical species belonging to D. sect. Drosera are divided into two lineages mostly defined by chromosome number. In the present work, a comprehensive taxonomic treatment for the Brazilian Drosera is presented, together with a revision of the previously poorly known Andean species, descriptions of new species and a molecular phylogeny focusing on the two Neotropical lineages of D. sect. Drosera. Although the phylogenetic analyses here performed did not present conclusive results regarding relationships at the species level, they shed light on the importance of some morphological traits for the classification of these species and back up the species circumscriptions adopted in recent works. Combined with data covering all the other lineages in a global phylogeny of Drosera, these results will be of crucial importance for the proposition of a new infrageneric classification of the genus. 1 Resumo Drosera é um gênero de plantas canívoras pertencente às Droseraceae e compreende cerca de 250 espécies distribuídas no mundo todo. U grande número de novas espécies e combinações foi publicado para o Brasil e os Neotrópicos nas últimas décadas, no entanto, ainda carecia-se de um compreensivo trabalho taxonômico lidando com tais espécies. Além disso, trabalhos filogenéticos anteriores focaram no entendimento das relações entre os grupos definidos por morfologia e distribuição geográfica, revelando que todas as classificações infragenéricas já propostas eram incongruentes com a filogenia e que as espécies Neotropicais pertencentes a D. sect. Drosera se dividiam em duas linhagens definidas, principalmente, por número cromossômico.
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