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Bru O Tomio Goto Taxo Omia De Glomeromycota: Revisão

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Bru O Tomio Goto Taxo Omia De Glomeromycota: Revisão BRUO TOMIO GOTO TAXOOMIA DE GLOMEROMYCOTA: REVISÃO MORFOLÓGICA, CHAVES DICOTÔMICAS E DESCRIÇÃO DE OVOS TÁXOS RECIFE 2009 II Universidade Federal de Pernambuco Centro de Ciências Biológicas Departamento de Micologia Pós-Graduação em Biologia de Fungos (ível Doutorado) TAXOOMIA DE GLOMEROMYCOTA: REVISÃO MORFOLÓGICA, CHAVES DICOTÔMICAS E DESCRIÇÃO DE OVOS TÁXOS 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 III 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 AGRADECIMETOS 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 SUMÁRIO Pág. Agradecimentos V Lista de Figuras X Lista de Tabelas XIV Resumo XV Abstract XVI Introdução Geral 17 Capítulo 1 – Revisão de Literatura 19 Micorrizas arbusculares - aspectos gerais 20 História da classificação dos FMA 20 Diversidade de Fungos Micorrízicos 32 omes e epítetos no filo Glomeromycota 35 Evolução dos nomes da estrutura reprodutiva básica “o esporo” 47 Aspectos morfológicos para identificação de FMA 49 Esporocarpo 52 Coloração dos esporocarpos 54 Perídio 55 TamanHo e forma dos esporocarpos 56 Hifas do plexo central do esporocarpo 57 Estrutura da parede 57 Tipos de camada da parede do esporo 60 Evanescente 60 Laminada 62 Unitária 63 Flexível 64 Amorfa 65 Expansiva 67 Germinativa 68 CHanfranulada 69 Grau de separação das camadas 69 Ornamentação da parede 70 Coloração dos glomerosporos 71 Goto, B.T. Taxonomia de Glomeromycota: revisão morfológica... 8 Efeito da resina sobre os glomerosporos 75 Dimensão dos glomerosporos 77 Forma do glomerosporo 79 Hifa de sustentação 79 úmero de Hifas por esporo 79 Sáculo esporífero 80 Bulbo suspensor 80 Estruturas especializadas de germinação (placas germinativas e orbs) 81 Reação em Melzer 88 Problemas de interpretação dos caracteres morfológicos 91 CHaves para identificação de FMA 93 Conceito de espécie no filo Glomeromycota 94 Material e Métodos 103 Capítulo 2 – Resultados e Discussão 106 Revisão dos caracteres morfológicos 109 Esporocarpos 109 Glomerosporos 113 Placa germinativa e orb 129 CHaves dicotômicas para gêneros e espécies de FMA 139 Descrições de algumas espécies de FMA 155 Conclusões 253 Perspectivas para a taxonomia do Filo Glomeromycota 253 Referências Bibliográfica 254 Glossário 269 Anexos 275 Goto, B.T. & Maia, L.C. 2005. Sporocarpic species of arbuscular mycorrhizal fungi (Glomeromycota), with a new report from Brazil. Acta Botanica Brasilica 19 (3): 633-637 276 Goto, B.T. & Maia, L.C. 2006. Glomerospores: a new denomination for the spores of Glomeromycota, a group molecularly distinct from the Zygomycota. Mycotaxon 96 : 129 - 132 281 Goto, B.T. & Maia, L.C. 2006. Contribution to the study of Endogonales in Brazil: the first record of Peridiospora tatachia . Mycotaxon 96 : 327 - 332. 285 Goto, B.T.; Moreira, M.; Tsai, S.M.; Cardoso, E.J.B.N.; Maia, L.C. 2008. Notas sobre Acaulospora bireticulata Rothwell & Trappe e primeiro registro de Acaulospora koskei Blaszk. para o Brasil. Acta Botanica Brasilica 22(2): 579- 291 583 Goto, B.T.; Maia, L.C & Oehl, F. Ambispora brasiliensis , a new ornamented Goto, B.T. Taxonomia de Glomeromycota: revisão morfológica... 9 species in the arbuscular mycorrhiza forming Glomeromycetes. Mycotaxon 105 : 11-18. 296 Goto, B.T.; Costa, C.M.C. & Maia, L.C. First record of Glomus halonatum Rose & Trappe (Glomeromycota) in South America - comments on the morphological characteristics of the species (Mycotaxon). 304 Goto, B.T.; & Maia, L.C. Pacispora velloziensis , a new species of arbuscular mycorrhiza-forming Glomeromycetes from Brazil (Mycotaxon) 308 Goto, B.T.; Mergulhão, A.C.E.S. & Maia, L.C. Notas sobre Acaulospora excavata Ingleby &Walker com revisão dos padrões de ornamentação em Acaulosporaceae. (Acta Botanica Brasilica). 312 Goto, B.T.; Maia, L.C & Oehl, F. Racocetra intraornata and Dentiscutata colliculosa , two new species in the Glomeromycetes from Northeastern Brazil with diagnostic spore wall ornamentations (Mycologia). 332 Goto, B.T.; Silva, G.A.; Trufem, S.F.B. & Maia, L.C. Glomus granulatum , a 356 new species of arbuscular mycorrhizal fungus from Brazil. (Mycotaxon) Goto, B.T. Taxonomia de Glomeromycota: revisão morfológica... 10 LISTA DE FIGURAS Pág. Figura 1 . Árvore filogenética com base em dados morfológicos e moleculares proposta por Morton & Redecker (2001). (Fonte: http://invam.caf.wvu.edu) 23 Figura 2 . Árvore filogenética mostrando um clado com os representantes dos fungos micorrízicos arbusculares definitivamente separados dos Zygomycetes (fonte: Schuessler et al. 2001). 25 Figura 3 . Detalhe das quatro ordens e das sete famílias que compõem o novo filo Glomeromycota (Fonte: Schuessler et al. 2001). 25 Figura 4. Árvore filogenética com inclusão da família Pacisporaceae (http://www.lrz-muenchen.de/~schuessler/amphylo/). 26 Figura 5 . Árvore filogenética mostrando a clado correspondente à família Ambisporaceae (Fonte: Walker et al. 2007). 30 Figura 6 – Tipos de parede ( sensu Walker 1983). G – Germinativa, P – Perídio, X – Expansivo, E – Evanescente, L – laminada, U – Unitária, M – Membranosa, C – Coriácea, A – Amorfa, R – Chafranulada. 50 Figura 7 A-G. Diversidade de esporocarpos em espécies de Glomus . A – Detalhe da organização dos esporos no esporocarpo de Gl. coremioides . B – Esporocarpo de Gl. clavisporum . C – Corte transversal de esporocarpos de Gl. cuneatum . D – Parte do esporocarpo de Gl. fulvum . E – Esporocarpos de Gl. aureum . F – Esporocarpo de Gl. antarcticum . G – Esporocarpo de Gl. glomerulatum . 112 Figura 8 A-F. Diversidade estrutural do perídio. A – Perídio de Gl. coremioides . B – Hifas ponteagudas no perídio de Glomus sp. C – Hifas em paliçada em Gl. liquidambaris . D – Perídio frouxamente arranjado em Gl. macrocarpum . E – Hifas sinuosas em Gl. sinuosum . F – Hifas entrelaçadas e coradas de Gl. cuneatum . 113 Figura 8. A -F Tipos de parede em glomerosporos. A- Laminada em Scutellospora sp1. B- Laminada expansiva em De biornata . C- Unitária em Scutellospora sp2. D- Flexível ou membranosa, em Ac. morrowiae. E- Evanescente expansiva, em Gl. halonatum . F- Amorfa, em Ac. foveata . 115 Figura 10 A-O. A- Acaulospora alpina . B- Ac. undulata . C- Ac. excavata . D- Ac. foveata . E- Ac. bireticulata . F- Acaulospora sp. G- Ac. denticulata . H- Ac. rehmii . I- Ac. tuberculata . J- Scutellospora biornata . K- Scutellospora sp1. L- Scutellospora sp2. M- Sc. gregaria . N- Sc. verrucosa . O- Glomus multiforum . 123 Figura 11 A-F. Variação na coloração dos glomerosporos. A- Ac. undulata . B- Gl. tortuosum. C- Gl. clavisporum . D- Gl. australe . E- Gl. constrictum . F- Gl. badium . 129 Figura 12 A-F. Diversidade de placas germinativas em glomerosporos gigasporóides. A – Placa germinativa complexa em De. biornata . B – Detalhe da
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