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The Fern Family Blechnaceae: Old and New

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The Fern Family Blechnaceae: Old and New ANDRÉ LUÍS DE GASPER THE FERN FAMILY BLECHNACEAE: OLD AND NEW GENERA RE-EVALUATED, USING MOLECULAR DATA Tese apresentada ao Programa de Pós-Graduação em Biologia Vegetal do Departamento de Botânica do Instituto de Ciências Biológicas da Universidade Federal de Minas Gerais, como requisito parcial à obtenção do título de Doutor em Biologia Vegetal. Área de Concentração Taxonomia vegetal BELO HORIZONTE – MG 2016 ANDRÉ LUÍS DE GASPER THE FERN FAMILY BLECHNACEAE: OLD AND NEW GENERA RE-EVALUATED, USING MOLECULAR DATA Tese apresentada ao Programa de Pós-Graduação em Biologia Vegetal do Departamento de Botânica do Instituto de Ciências Biológicas da Universidade Federal de Minas Gerais, como requisito parcial à obtenção do título de Doutor em Biologia Vegetal. Área de Concentração Taxonomia Vegetal Orientador: Prof. Dr. Alexandre Salino Universidade Federal de Minas Gerais Coorientador: Prof. Dr. Vinícius Antonio de Oliveira Dittrich Universidade Federal de Juiz de Fora BELO HORIZONTE – MG 2016 Gasper, André Luís. 043 Thefern family blechnaceae : old and new genera re- evaluated, using molecular data [manuscrito] / André Luís Gasper. – 2016. 160 f. : il. ; 29,5 cm. Orientador: Alexandre Salino. Co-orientador: Vinícius Antonio de Oliveira Dittrich. Tese (doutorado) – Universidade Federal de Minas Gerais, Departamento de Botânica. 1. Filogenia - Teses. 2. Samambaia – Teses. 3. RbcL. 4. Rps4. 5. Trnl. 5. TrnF. 6. Biologia vegetal - Teses. I. Salino, Alexandre. II. Dittrich, Vinícius Antônio de Oliveira. III. Universidade Federal de Minas Gerais. Departamento de Botânica. IV. Título. À Sabrina, meus pais e a vida, que não se contém! À Lucia Sevegnani, que não pode ver esta obra concluída, mas que sempre foi motivo de inspiração. Lucia, você faz muita falta! “A verdade pertence a todos e a ninguém. É propriedade pública e efêmera, ou não é o tipo de verdade que chamamos de ciência. Mas o erro é eterno porque, tendo ocorrido uma vez, existe, como uma família infeliz, à sua própria maneira.” A medida de todas as coisas. Ken Alder AGRADECIMENTOS - À minha família, pelo incondicional apoio desde a graduação e por acreditar no sonho; - À Sabrina, que se tornou família durante a graduação, e permanece mesmo nas ausências; - Ao meu orientador, Alexandre Salino, pela orientação, pelo apoio e discussões, mesmo a distância, e por ter me acolhido lá em 2009, o ainda desconhecido sulista em sua casa; - Ao meu coorientador, Vinícius Antonio de Oliveira Dittrich, por todas as discussões e por abraçar este projeto!; - Ao Dr. Alan Smith, que sem sua ajuda, aquela olhada na planta de herbário, e sapiência milenar, esta tese ainda estaria patinando nos Blechna! - Ao Raymond Cranfill, que de seus rascunhos guardados a 15 anos numa gaveta, mostrou um caminho a seguir; - Aos professores do PGBOT, pela convivência, mesmo que em curtos períodos, mas que abriram a mente a novas ideias e oportunidades, meus mais sinceros agradecimentos; - Aos pteridólogos de plantão, pela vivência e conhecimento compartilhado: Thais, Fran, Raquel, Rozi, Fifão, Jeferson, Neto, Lili, Juliana; - Aos amigos mineiros, ou nem tanto mineiros como eu, em ordem aleatória, seja pelo cantinho cedido na república e pela amizade construída: Ana, Eric, Leila, Rozi, Fifão, Thais, Leandro, Marcelo Bueno, Vanessa P., Maria José, Bruno, Aline, Cleber, Barbara, Vanessa R, Fernanda; - Aos amigos de graduação da FURB, que até hoje se divertem e riem juntos: Chico, Roger, Sabrina, Marcos e Durda; - Ao CNPq pela aprovação do projeto no Edital Universal, que possibilitou as viagens de coleta e o financiamento dos experimentos; - Ao Programa de Bolsas do Fundo de Apoio à Manutenção e ao Desenvolvimento da Educação Superior – FUMDES, de Santa Catarina, que ao conceder a bolsa de estudos, permitiram as viagens do doutorado no eixo Blumenau-Belo Horizonte; - Ao Dr. Daniel Tejeto Diez do México e ao Christian Adolfo López do Panamá, pela cia e pelas coletas nos dois países! - Aqueles que doaram amostras, dentre eles: Michel Kessler, Ebihara, Ralf Kanpp, Funez, dentre tantos outros, bem como ao Jardim Botânico de Paris, ao Missouri Botanical Garden e ao Kew pelas amostras; elas foram fundamentais; - À FURB, onde construí e construo minha carreira, sem a qual não teria chego onde estou, especialmente pelo afastamento nos últimos seis meses para conclusão da tese; - Aos amigos docentes, discentes e tantos outros, dentre eles do Lab. de Botânica: Sidney, Roberta, Morilo, Felipe, Karl, Hanna, Fábio, Deizi, Guilherme, Nando, Giesta; do Departamento de Ciências Naturais: Alessandra, Simone, Giasson, Hercilio, Zelinda e tantos outros, pela vivência e convivência! - Àqueles que ajudaram nas discussões, desesperos, descobertas e leituras: Aline, Thais, Karina Proite, João Batista; - Ao IFFSC, projeto do qual faço parte desde 2005, do qual tenho imenso orgulho e toda a sua equipe, em especial ao Alexandre Vibrans, que desde 2007 se tornou um parceiro de trabalho; - Aos antigos mestres pelos primeiros passos acompanhados, dentre eles: Karin, Uhlmann, Juarez, Bacca, Sidney, Lucia, Rosete, Rudi e Zelinda. ÍNDICE RESUMO GERAL............................................................................................................. i ABSTRACT....................................................................................................................... ii INTRODUÇÃO GERAL................................................................................................... 1 CAPÍTULO 1. The fern family Blechnaceae: old and new genera re-evaluated, using molecular data..................................................................................................................... 6 Abstract................................................................................................................... 7 Introduction............................................................................................................. 8 Material and methods............................................................................................ 11 Results.................................................................................................................... 13 Discussion.............................................................................................................. 14 Conclusion............................................................................................................. 29 Acknowledgments................................................................................................. 30 References……………….…………………………………………...……...….. 30 Appendix…………………………………………………..………………......... 43 CAPÍTULO 2. A classification of Blechnaceae: New genera, resurrected names, and combinations…………………………..……………………………….………….…...... 63 Abstract…………….………………………….……………….……………..... 64 Introduction……………..………………………..……..……….……….…….... 65 Methods………………………………………………………….……….……… 68 Results…………………………………………………………….…….……..… 69 Characters…………………………………...……....…………….…….……….. 69 Taxonomic treatment………….……………………..………....….….……….… 77 Key to the Blechnaceae genera………………………..…...………..………...… 78 Acknowledgments…………….......……………………….....................……... 129 References…………………………....………….....................…………..…… 130 CONCLUSÕES GERAIS………...………..................……………………………… 147 REFERÊNCIAS GERAIS…………………………....................…………………… 148 LISTA DE FIGURAS CAPÍTULO 1. Figure 1. Majority-rule consensus phylogram from Bayesian Inference analysis from concatenated dataset, for three chloroplast regions (rbcL, rps4, trnL-trnF), showing phylogenetic hypothesis for relationship among Blechnaceae taxa. Bayesian Inference posterior probabilities, Maximum Likelihood bootstrap values and Maximum Parsimony bootstrap values (%) are indicated, respectively, on the branches. Thick lines indicate strong support (posterior probability = 1.00 and maximum likelihood bootstrap support and maximum parsimony = 100%)……………………………………...…………………….. 57 CAPÍTULO 2. FIGURE 1. General appearance of the three genera of Blechnaceae subfamily Woodwardioideae. A: Anchistea virginica (P01513597); B: Lorinseria areolata (P01509531), C: Woodwardia spinulosa (P01557309). Vouchers are indicated by herbarium barcodes. Herbarium abbreviations according Thiers (2015)……………….……………141 FIGURE 2. General appearance of the three genera of Blechnaceae subfamily Stenochlaenoideae. A: Salpichlaena volubilis (MO3588318); B: Stenochlaena scandens (P01420182); C: Telmatoblechnum serrulatum (MO2003917). Vouchers are indicated by herbarium barcodes. Herbarium abbreviations according Thiers (2015)…………………141 FIGURE 3. General appearance of the genera of Blechnaceae subfamily Blechnoideae (A- D). A: Austroblechnum penna-marina (FURB06706); B: Blechnidium melanopus (P02439094); C: Blechnopsis orientalis (P01406555); D: Blechnum occidentale (FURB06701); E: Brainea insignis (P01482127); F: Cleistoblechnum eburneum (P00627565); G: Cranfillia fluviatilis (P01606472); H: Diploblechnum fraseri (P01557366); I: Doodia aspera (P01618184). Vouchers are indicated by herbarium barcodes. Herbarium abbreviations according Thiers (2015)…………………………………………………...142 FIGURE 4. General appearance of the genera of Blechnaceae subfamily Blechnoideae (G- S). A: Icarus filiformis (WELT P009478); B: Lomaria nuda (P01557820); C: Lomaridium fragile (MEXU 149093); D: Lomariocycas aurata (QCA7004842); E: Neoblechnum brasiliense (FURB06585); F: Oceanopteris cartilaginea (P01630912). G: Parablechnum cordatum (FURB06639); H: Sadleria pallida (MEXU 956078); I Struthiopteris spicant (P01658104); I:. Vouchers are
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