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TESE Victor Rafael Matos Coimbra.Pdf UNIVERSIDADE FEDERAL DE PERNAMBUCO CENTRO DE BIOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA DE FUNGOS VICTOR RAFAEL MATOS COIMBRA RIQUEZA E ASPECTOS MOLECULARES DE Gymnopus (OMPHALOTACEAE, AGARICALES) NO NORTE E NORDESTE BRASILEIRO Recife 2017 VICTOR RAFAEL MATOS COIMBRA RIQUEZA E ASPECTOS MOLECULARES DE Gymnopus (OMPHALOTACEAE, AGARICALES) NO NORTE E NORDESTE BRASILEIRO Tese apresentada ao Programa de Pós- Graduação em Biologia de Fungos, Área de Concentração Micologia Básica, da Universidade Federal de Pernambuco como requisito parcial para a obtenção do título de doutor em Biologia de Fungos. Orientador: Profa. Dra. Tatiana Baptista Gibertoni. Recife 2017 Catalogação na fonte Elaine Barroso CRB 1728 Coimbra, Victor Rafael Matos Riqueza e aspectos moleculares de Gymnopus (Omphalotaceae, Agaricales) do Norte e Nordeste brasileiro / Victor Rafael Matos Coimbra - 2017. 103 folhas: il., fig., tab. Orientadora: Tatiana Baptista Gibertoni Tese (doutorado) – Universidade Federal de Pernambuco. Centro de Biociências. Programa de Pós-Graduação em Biologia de Fungos, Recife, 2017. Inclui referências e apêndices 1. Agaricus (cogumelo) 2. Filogenia 3. Mata Atlântica I. Gibertoni, Tatiana Baptista (orient.) II. Título 579.59 CDD (22.ed.) UFPE/CB-2018-034 VICTOR RAFAEL MATOS COIMBRA RIQUEZA E ASPECTOS MOLECULARES DE Gymnopus (OMPHALOTACEAE, AGARICALES) NO NORTE E NORDESTE BRASILEIRO Tese apresentada ao Programa de Pós- Graduação em Biologia de Fungos, Área de Concentração Micologia Básica, da Universidade Federal de Pernambuco, como requisito parcial para obtenção do título de doutor em Biologia de Fungos. Aprovada em: 21/02/2017 COMISSÃO EXAMINADORA ___________________________________________ Dra. Tatiana Baptista Gibertoni / Universidade Federal de Pernambuco ___________________________________________ Dra. Bianca Denise Barbosa da Silva / Universidade Federal da Bahia __________________________________________ Dr. Rafael Batista Louzada / Universidade Federal de Pernambuco __________________________________________ Dr. Gladstone Alves da Silva / Universidade Federal de Pernambuco __________________________________________ Dra. Laise de Holanda Cavalcanti Andrade / Universidade Federal de Pernambuco AGRADECIMENTOS Primeiramente, agradeço a Deus por me proporcionar a realização de diversos sonhos pessoais e profissionais durante o desenvolvimento desse doutorado. Esses últimos quatro anos foram uma verdadeira benção, se transformando em um capítulo especial de minha história de vida e de meu currículo acadêmico. Agradeço também à toda minha família, especialmente a minha mãe (Beta), meus irmãos (Eduardo e Ana), minha tia Sandra e meus avôs (Adalberto e Júlia), por todo o amor e cuidado devotados a mim. Eles foram e continuam sendo alicerces fundamentais na minha vida. Extendo meus agradecimentos também a minha namorada (Kaisa Nascimento), minha grande “fã” e incentivadora. Meus mais sinceros agradecimentos a meus orientadores Dra. Tatiana B. Gibertoni (UFPE) e Dr. Felipe Wartchow (UFPB). O apoio e suporte de vocês foram essenciais para o desenvolvimento dessa pesquisa e para minha formação como cientista. A todos os meus colegas de laboratório da UFPE e da UFPB, agradeço pela companhia e amizade durante todo esse tempo. Extendo os agradecimentos aos demais colegas de doutorado e do Departamento de Micologia da UFPE. Agradeço também a todos membros do Instituto de Biologia e Meio Ambiente (BioEnv - Institutionen för biologi och miljövetenskap) da Universidade de Gotemburgo, Suécia, pelo suporte para o desenvolvimento de parte de minha pesquisa. Um agradecimento especial é destinado a Dr. Ellen Larsson (orientadora no exterior) e Dr. Rolf Henrik Nilsson (colaborador). Agradeço também pela calorosa acolhida e pelos ensinamentos transmitidos por todos os membros da Sociedade Sueca de Micologia (SMF – Sveriges Mykologiska Förening), dentre eles Sten Svantesson, Kill Persson, Mikael Jeppson e Stig Jacobsson. A todos os micólogos que contribuíram diretamente ou inidiretamente nesse projeto, ou mesmo em projetos paralelos e na minha formação intelectual, meus mais sinceros agradecimentos. Dentre eles, destaco Dr. Gladstone Silva (UFPE, Brasil), Dr. Vladimír Antonín (Moravské Zemské Muzeum, Rep. Tcheca), Dr. Pierre-Arthur Moreau (Université du Droit et de la Santé Lille 2, França), Dr. Michal Tomsovský (Mendel University, Rep. Tcheca), Dr. Timothy Baroni (SUNY Cortland, Estados Unidos) e Dr. Karl-Henrik Larsson (University of Oslo, Noruega). Meus mais sinceros agradecimentos a todos os órgãos e instituições que financiaram direta ou indiretamente esse projeto: FACEPE – Fundação de Amparo a Ciência e Tecnologia de Pernambuco (IBPG 0827-2.03/12, AMD 0218-2.00/13, APQ 0788-2.03/12), CNPQ – Conselho Nacional de Desenvolvimento Científico e Tecnológico (SWE 232695/2014-8, Sisbiota 563342/ 2010-2, PROTAX 562106/2010-3) e CAPES - Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES/SIU 008/13). RESUMO Com o intuito de contribuir com o conhecimento taxonômico e filogenético de espécies do gênero Gymnopus (Agaricales, Basidiomycota) ocorrentes em áreas de Mata Atlântica e Floresta Amazônica do Norte e Nordeste do Brasil, foram realizadas 47 expedições (22 localidades em nove estados) de coleta, entre os anos de 2012 e 2015. Foram coletadas ao todo 104 amostras (≅ 700 basidiomas), sendo posteriormente descartadas 21 delas. As 83 amostras restantes corresponderam a 25 espécies, sendo 15 delas novas para a ciência, uma proposta como nova combinação e três registradas pela primeira vez para o Brasil. Desta forma, houve um incremento na riqueza de Gymnopus no Brasil de aproximadamente 95% (de 19 para 37 espécies). Dentre as cinco seções reconhecidas na sistemática atual do gênero, apenas membros de Gymnopus seção Impudicae (quatro espécies) e Gymnopus seção Vestipedes (21 espécies) foram coletados nas regiões estudadas. Das 88 amostras submetidas aos procedimentos de biologia molecular, 66 delas obtiveram sucesso (75%), resultando em 116 sequências de DNA de boa qualidade (62 de ITS, 47 de LSU e sete de TEF1-ɑ). Baseado exclusivamente em análises morfológicas, foi possível confirmar as identidades de G. collybioides, G. coracicolor e G. subpruinosus, enquanto, as análises polifásicas (morfológicas e filogenéticas multigênicas - ITS+LSU+ TEF1-ɑ) deram suporte a determinações de 15 novos táxons para a ciência, bem como a identificação de G. albelloavellaneus, G. alnicolus, G. gibbosus, G. montagnei, G. omphalodes, G. polygrammus e G. subcyathiformis. Palavras-chave: Cogumelos. Taxonomia. Filogenia. Floresta Amazônica. Mata Atlântica. ABSTRACT With the aim of contributing to the taxonomic and phylogenetic knowledge of species of the genus Gymnopus (Agaricales, Basidiomycota) occurring in Atlantic and Amazon Forest areas of Brazil’s North and Northeast, 47 collection expeditions (22 sites in nine states) were carried out, between the years 2012 and 2015. A total of 104 samples (≅ 700 basidiomas) were collected, being 21 of them later discarded. The 83 remaining samples corresponded to 25 species, being 15 of them new to science, one proposed as new combination and three recorded for the first time to Brazil. Thus, there was an increase in the Gymnopus’ richness in Brazil of about 95% (from 19 to 37 species). Among the five sections recognized in the current systematics of the genus, only members of Gymnopus section Impudicae (four species) and Gymnopus section Vestipedes (21 species) were collected in the studied regions. Of 88 samples submitted to molecular biology procedures, 66 of them were successful (75%), resulting in 116 good quality DNA sequences (62 of ITS, 47 of LSU and seven of TEF1-ɑ). Based exclusively on morphological analyses, it was possible to confirm the identities of G. collybioides, G. coracicolor and G. subpruinosus, while the polyphasic analyses (morphological and multi-gene - ITS+LSU+ TEF1-ɑ) gave support to the determinations of 15 new taxa to science as well as the identification of G. albelloavellaneus, G. alnicolus, G. gibbosus, G. montagnei, G. omphalodes, G. polygrammus and G. subcyathiformis. Key-words: Mushrooms. Taxonomy. Phylogeny. Amazon Forest. Atlantic Forest. LISTA DE ABREVIATURAS E SIGLAS CTAB Brometo de Cetil Trimetilamonio. FLONA Floresta Nacional. FLOTA Floresta Estadual. ITS DNAr Espaçadores transcritos internos 1 e 2, que circundam o gene 5.8S e são flanqueados pelos genes SSU (18S) e LSU (28S). LSU DNAr Gene da subunidade maior do ribossomo, também conhecida como 28S. PCR Reação em cadeia da polimerase. REBIO Reserva Biológica. RESEX Reserva Extrativista. RPPN Reserva Particular do Patrimônio Natural. TEF1-ɑ Subunidade 1 alpha do fator de elongamento de tradução. UFPA Universidade Federal do Pará. UFPE Universidade Federal de Pernambuco. UFPB Universidade Federal da Paraíba. SUMÁRIO 1 INTRODUÇÃO....................................................................................................................14 2 FUNDAMENTAÇÃO TEÓRICA......................................................................................16 2.1 Sistemática de Gymnopus...................................................................................................16 2.2 Diversidade no Brasil e no mundo......................................................................................20 2.3 Importância ecológica e econômica....................................................................................21 3 MATERIAIS E MÉTODOS...............................................................................................23
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