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Bruno Tomio Goto Taxonomia De Glomeromycota

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Bruno Tomio Goto Taxonomia De Glomeromycota BRUNO TOMIO GOTO TAXONOMIA DE GLOMEROMYCOTA: REVISÃO MORFOLÓGICA, CHAVES DICOTÔMICAS E DESCRIÇÃO DE NOVOS TÁXONS RECIFE 2009 II BRUNO TOMIO GOTO TAXONOMIA DE GLOMEROMYCOTA: REVISÃO MORFOLÓGICA, CHAVES DICOTÔMICAS E DESCRIÇÃO DE NOVOS TÁXONS Tese apresentada ao Curso de Pós- Graduação em Biologia de Fungos da Universidade Federal de Pernambuco, como parte das exigências para obtenção do grau de Doutor. Área de concentração: Micologia Básica Orientadora: Dra. Leonor Costa Maia Co-orientador: Dr. Gladstone Alves da Silva Recife Fevereiro/2009 Goto, B.T. Taxonomia de Glomeromycota: revisão morfológica... 358 Goto, Bruno Tomio Taxonomia de Glomeromycota: revisão morfológica, chaves dicotômicas e descrição de novos táxons / Bruno Tomio Goto. – Recife: O Autor, 2009. vi, 357 folhas : il., fig., tab. Tese (doutorado) – Universidade Federal de Pernambuco. CCB. Biologia de Fungos, 2009. Inclui bibliografia, anexo e glossário. 1. Glomeromycota – Taxonomia 2. Fungos micorrízicos arbusculares 3. Morfologia I Título. 582.281.2 CDU (2.ed.) UFPE 579.53 CDD (22.ed.) CCB – 2009-136 III BRUNO TOMIO GOTO TAXONOMIA DE GLOMEROMYCOTA: REVISÃO MORFOLÓGICA, CHAVES DICOTÔMICAS E DESCRIÇÃO DE NOVOS TÁXONS Tese defendida e aprovada pela banca examinadora Aprovada em 11/02/2009 BANCA EXAMINADORA Dra. Leonor Costa Maia – (UFPE) Dra. Adriana Mayumi Yano Melo – (UNIVASF) Dra. Rosilaine Carrenho – (UEM) Dra. Sandra Farto Botelho Trufem (Universidade de São Carlos) Dr. Francisco Adriano de Souza (Embrapa-Agrobiologia) IV Aos meus Pais Chouji Goto e Amilce Goto. À Esposa Rebeca Garrett e Filha Júlia Goto. Aos verdadeiros mestres Leonor Maia, Sandra Trufem, Maria Auxiliadora, José Luiz Bezerra e Fritz Oehl. Dedico e ofereço V AGRADECIMENTOS Agradeço À Profa. Sandra Farto Botelho Trufem pela paciência e disponibilidade em ensinar-me os primeiros passos na taxonomia desse grupo de fungos; À Profa. Maria Auxiliadora de Queiroz Cavalcanti pelos ensinamentos, estímulos e disposição em ensinar taxonomia de fungos; À Profa. Adriana Mayumi Yano Melo por ter me iniciado nas pesquisas em taxonomia; Ao Prof. Gladstone Alves da Silva, co-orientador, consultor e amigo pela disponibilidade em ajudar e discutir os tópicos da tese; À Profa. Uided Maaze Tiburcio Cavalcanti pelos ensinamentos e principalmente pela paciência no dia-a-dia do laboratório; À Profa. Laize de Holanda Cavalcanti pelo estímulo ao projeto de tese e principalmente nos conselhos para a formulação das chaves de identificação; Ao Prof. José Luiz Bezerra pelos ensinamentos e pelo envio de materiais importantes para o desenvolvimento da tese; Ao Prof. Fritz Oehl pela disposição, paciência para me ensinar taxonomia de FMA e principalmente pelas dícas e vislumbres nesse interessante grupo de fungos; À Dra. Renata Gomes de Souza pela ajuda, paciência e principalmente pela disposição em fazer taxonomia de FMA; Aos colegas Angelo, Nicácio, Moacir, Araeska, Heloísa, Camila, João, Thiago, Vilma, Indra, Érika, Cláudia do laboratório de micorrizas pela ajuda e momentos de descontração; Aos colegas Ricardo, Paul, Felipe, Jadergudson e Allyne do laboratório de taxonomia de fungos pela amizade e ensinamentos em taxonomia dos outros grupos de fungos que contribuíram bastante para a construção de meu conhecimento; À Profa. Elaine Malosso pelos momentos de inspiração para encontrar encantamento nos estudos moleculares; Aos pesquisadores Vagner Cortéz, Luiz Gusmão; Iuri Baseia, Ricardo Berbara, Francisco de Souza, Rosilaine Carrelho, Sidney Stürmer, Milene Moreira, Camila Nobre e Natoniel Franklin de Melo pelo envio de material biológico, incentivo e discussões valiosas durante o meu amadurecimento científico; VI Aos meus pais Chouji Goto, Amilce Goto e ao irmão Teruo Goto pelo companheirismo e principalmente pelos estímulos durante a caminhada acadêmica; Aos meus amigos Joaquim Otto, Diego Diêgues, Isabela Alcantara, Regina Melo, Reginaldo Lima Neto pela paciência devido a minha ausência durante o desenvolvimento da tese; A minha esposa Rebeca Garrett e filha Júlia Goto por compreenderem a minha ausência durante a minha jornada acadêmica. E finalmente à CAPES pela bolsa de doutorado e a Pós-Graduação em Biologia de Fungos. Goto, B.T. Taxonomia de Glomeromycota: revisão morfológica... 7 RESUMO GERAL A taxonomia dos fungos micorrízicos arbusculares (FMA) vem sofrendo grandes alterações durante os últimos 20 anos e isso é facilmente evidenciado pelas mudanças na classificação desses fungos que deixaram de ser meros representantes da família Endogonaceae, passando para Ordem Glomales, ainda dentro dos Zygomycetes, e finalmente culminando no filo Glomeromycota onde foram definitivamente segregados dos Zygomycota. Todas essas alterações na classificação estão relacionadas às mudanças na interpretação agregada aos caracteres relacionados exclusivamente aos glomerosporos onde toda a taxonomia morfológica está embasada e também aos dados moleculares. Essas mudanças têm sido incorporadas à taxonomia do grupo, com as descrições de novas famílias, gêneros e espécies que incluíram evidências morfológicas adicionais já dentro do novo filo. Apesar dessas alterações recentes, nenhuma análise morfológica detalhada foi realizada desde a proposta do filo Glomeromycota, o que torna a taxonomia do grupo mais frágil considerando que a maior parte das propostas de novas famílias e gêneros estão baseadas quase que exclusivamente em dados moleculares. Por essa razão, o objetivo deste trabalho foi revisar os caracteres morfológicos com base em evidências disponíveis na literatura assim como nas espécies herborizadas, mantidas em coleções de germosplasma e principalmente em esporos de campo onde a maior parte das espécies tem sido descritas com intuito de fornecer análises detalhadas desses caracteres usados para a identificação, incorporar à taxonomia do grupo novas abordagens morfológicas e elaborar chaves para facilitar a identificação de gêneros e espécies de Glomeromycota. Para isso, todas as descrições de espécies foram analisadas desde a descrição de Glomus macrocarpum e Gl. microcarpum em 1844. Além disso, glomerosporos provenientes de coleções de cultura viva (EMBRAPA, UFPE, UFRRJ) e herborizadas (FL Herbarium, FH, Universidad de Havanna, Universidad de Buenos Aires e material da coleção pessoal do prof. Fritz Oehl) foram avaliadas. A principal fonte de dados foram os glomerosporos provenientes de amostras de campo de diversos ecossistemas brasileiros e alguns da América Latina. Glomerosporos provenientes de amostras de campo foram extraídos do solo por peneiramento úmido seguido de centrifugação em água e sacarose a 40%. Após a extração os glomerosporos foram separados por cor e tamanho e montados entre lamina e lamínula com PVLG e PVLG + Reagente de Melzer (1:1 v/v) para posterior análise em microscópio composto. Culturas armadilhas com solo nativo e Sorghum bicolor (L.) foram mantidas por três meses em média para captura das espécies de FMA. No total foram avaliadas 106 espécies de FMA pertencentes a todos os gêneros exceto Otospora, incluindo cinco novas espécies para a ciência, abrangendo todos os modos de desenvolvimento encontrado no filo Glomeromycota. Os caracteres morfológicos avaliados foram: cor, forma, tamanho dos esporocarpos, perídio e plexo central (cor, tamanho e forma); cor, tamanho, forma e modo de desenvolvimento dos glomerosporos; espessura, tipo e estrutura da parede, reação ao Melzer ou a resina (PVLG) e as estruturas especializadas de germinação (orbs e placas germinativas). Conclui-se que o único caráter morfológico compartilhado entre todos os representantes do filo Glomeromycota é a estrutura da parede, sendo assim os caracteres morfológicos considerados primários (mais úteis para a identificação das espécies de FMA) são: estrutura da parede, tipo de camada e ornamentação enquanto cor, forma, tamanho e reação ao Melzer ou resina são considerados secundários para a identificação de espécies. Palavras-chave: Glomeromycetes, taxonomia, FMA Goto, B.T. Taxonomia de Glomeromycota: revisão morfológica... 8 ABSTRACT The taxonomy of the arbuscular mycorrhizal fungi (AMF) has undergone major changes over the past 20 years and this is easily evidenced by changes in the classification of these fungi that ceased to be mere representatives of the family Endogonaceae, to Order Glomales, even within the Zygomycetes, and finally culminating in the phylum Glomeromycota where they were finally separated the Zygomycota. All these changes in classification are related to changes in interpretation of aggregate characters related only to glomerosporos where the whole is based morphological taxonomy and molecular data. These changes have been incorporated into the taxonomy of the group, with descriptions of new families, genera and species that included additional morphological evidence already in the new phylum. Despite these recent changes, no detailed morphological analysis was performed since the proposal of the phylum Glomeromycota, which makes the taxonomy of the group more fragile whereas most of the proposals of new families and genera are based almost exclusively on molecular data. Therefore, the objective of this study was to review the morphological characters based on evidence available in literature and on herbarium species held in collections of spores germosplasma and mainly in the field where most of the species have been described in order to provide Detailed analysis of these characters for the identification, taxonomy of the group to incorporate new approaches and develop morphological keys to facilitate the identification
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