Poaceae, Panicoideae), Como Subsídio À Sua Taxonomia

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Poaceae, Panicoideae), Como Subsídio À Sua Taxonomia LUIZ ANTONIO DE JESUS JUNIOR ESTUDOS MORFOANATÔMICOS NO COMPLEXO ICHNANTHUS PALLENS (SW.) MUNRO EX BENTH. (POACEAE, PANICOIDEAE), COMO SUBSÍDIO À SUA TAXONOMIA FEIRA DE SANTANA - BA 2013 UNIVERSIDADE ESTADUAL DE FEIRA DE SANTANA DEPARTAMENTO DE CIÊNCIAS BIOLÓGICAS PROGRAMA DE PÓS-GRADUAÇÃO EM BOTÂNICA ESTUDOS MORFOANATÔMICOS NO COMPLEXO ICHNANTHUS PALLENS (SW.) MUNRO EX BENTH. (POACEAE, PANICOIDEAE), COMO SUBSÍDIO À SUA TAXONOMIA LUIZ ANTONIO DE JESUS JUNIOR Dissertação apresentada ao Programa de Pós- graduação em Botânica da Universidade Estadual de Feira de Santana, como parte dos requisitos para obtenção do título de Mestre em Botânica. ORIENTADORA: PROFa. DRA. REYJANE PATRÍCIA DE OLIVEIRA CO-ORIENTADORA: PROFa. DRA. KELLY REGINA BATISTA LEITE FEIRA DE SANTANA - BA 2013 BANCA EXAMINADORA ___________________________________________________ Prof. Dr. Francisco de Assis Ribeiro dos Santos Departamento de Ciências Biológicas, UEFS ___________________________________________________ Profa. Dra. Bárbara Rosemar Nascimento de Araújo União Metropolitana para o Desenvolvimento da Educação e Cultura - UNIME ___________________________________________________ Profa. Dra. Reyjane Patrícia de Oliveira Departamento de Ciências Biológicas, UEFS Orientadora e Presidente da banca FEIRA DE SANTANA - BA 2013 À minha mãe e minha irmã, alicerces da minha vida, pelo apoio incondicional para alcançar meus objetivos. i AGRADECIMENTOS Agradeço sinceramente à minha orientadora Profa Dra. Reyjane Patrícia de Oliveira, pela oportunidade, amizade, paciência, compreensão e extrema dedicação, inclusive em seus dias de férias. À minha co-orientadora Profa Dra. Kelly Regina Batista Leite, pela amizade, apoio, dedicação e pelas horas gastas em videoconferências fora do seu horário de trabalho. À Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES), pela bolsa de mestrado concedida. À Universidade Estadual de Feira de Santana pela viabilização da infraestrutura para desenvolvimento deste trabalho. A todos os professores do Programa de Pós-graduação em Botânica, pela grande contribuição para minha formação. À Profa Dra. Claudia Elena Carneiro, coordenadora do Laboratório de Micromorfologia Vegetal (LAMIV), por disponibilizar as dependências deste para o desenvolvimento dos estudos anatômicos. Agradeço aos colegas do LAMIV pelo apoio, amizade e pelos momentos de descontração. Em especial ao Dr. Marcos da Costa Dórea e ao Dr. Paulino Pereira Oliveira, por sempre se mostrarem prestativos quando algum auxilio era solicitado. Ao Dr. Luiz Fernando Pascholati Gusmão, por disponibilizar o uso do microscópio com câmera acoplada para obtenção das fotomicrografias. À Fundação Oswaldo Cruz, por disponibilizar o uso do microscópio eletrônico de varredura. Aos funcionários do LABIO, pela amizade e por sempre se mostrarem prestativos. ii Ao Christian da Silva, pela grande amizade, pelo subsídio nas coletas dos materiais e constante apoio. Aos demais irmãos que trabalham com Poaceae: Tutti, Karena e Fabrício pelo auxílio constante. Às amigas Isys Mascarenhas e Kamilla Lopes pela ajuda na obtenção de materiais herborizados e pela grande amizade construída. A todos meus amigos da pós-graduação, pela troca de conhecimento e momentos de descontração proporcionados durante esse período. Em especial a: Aline Quaresma, Ariane, Dani Mendes, Diego Generoso, Gabizão, Grilo, Karol, Lara, Marcelo, Marloca, Nine, Priscila, Rodolfo, Seu Marlon, Snakinha e Taiara. Ao amigo Igor Monteiro, pela ajuda na confecção dos mapas e pelo companheirismo construído através de anos de parceria, obrigado irmão. A toda minha família, pelo incentivo, em especial à minha tia Madalena, pelo exemplo de pessoa. Agradeço à minha namorada Silvana Santos da Silva, pelas alegrias proporcionadas, pelo cuidado que tem comigo e pelo amor que me confia. Agradeço por estar todo esse tempo ao meu lado, apoiando as minhas decisões, independente de quais sejam! Eu te amo minha gata! À minha mãe, pelo amor incondicional, sempre me orientando com sabedoria e me estimulando a alcançar meus objetivos. Agradeço também a minha irmã, por sempre me apoiar e servir de exemplo de como enfrentar os osbstáculos da vida. iii RESUMO O gênero Ichnanthus P. Beauv. pertence à tribo Paspaleae, subfamília Panicoideae (Poaceae). Inclui atualmente ca. 33 espécies ocorrentes predominantemente em ambientes florestais, em especial na região Neotropical. Tem sido tradicionalmente reconhecido pelas lâminas foliares com base geralmente assimétrica, inflorescências paniculadas, espiguetas comprimidas lateralmente e apêndices aliformes na base do lema superior, cicatrizes desses apêndices ou apenas protuberâncias convexas na mesma posição. Uma das suas espécies, I. pallens (Sw.) Munro ex Benth., com cicatrizes na base do lema superior, inclui duas variedades (I. pallens var. pallens e I. pallens var. major (Nees) Stieber). Essa demonstra grande variação nos caracteres vegetativos e reprodutivos, com vários morfotipos cujas morfologias às vezes se sobrepõem, entre si e com outras espécies relacionadas, como I. tenuis (J.S Presl) Hitchc. e I. ruprechtii Döll, constituindo o complexo I. pallens. Com base em amostras obtidas em campo e em materiais depositados em herbários, foram realizadas análises morfoanatômicas envolvendo folhas e inflorescências em representantes desse complexo, a fim de verificar a ocorrência de variações inter e intraespecíficas, e se tais caracteres podem ser úteis para fins taxonômicos. Para as análises em secção transversal das lâminas foliares e dos pedúnculos das inflorescências, as amostras foram emblocadas em resina histológica e analisadas através de microscopia óptica. Para o estudo da superfície da epiderme foliar e dos antécios, foi utilizada, além da microscopia óptica, análises de microscopia eletrônica de varredura. Lâminas foliares e inflorescências também foram analisadas em microscópio estereoscópico. Foi observada uma forte distinção na anatomia foliar de I. pallens em relação às duas espécies analisadas, sendo designados três padrões principais envolvendo iv essa espécie: o „padrão I. pallens sensu stricto’, formado por I. pallens var. major e I. pallens var. pallens modos axillaris, delicado e sensu stricto; o „padrão I. pallens modo confertus‟, formado exclusivamente por I. pallens var. pallens modo confertus e „padrão I. pallens modo areolatus‟, composto apenas por I. pallens var. pallens modo areolatus. As duas outras espécies analisadas também representam padrões distintos (I. tenuis e I. ruprechtii). Os dados da anatomia do pedúnculo da inflorescência confirmam esses padrões, enquanto os caracteres macromorfológicos das inflorescências e da superfície do antécio em MEV apresentam muita sobreposição. Os caracteres morfoanatômicos avaliados no complexo I. pallens, tanto em níveis inter quanto intraespecíficos, possuem variação pouco significativa relacionada à distribuição geográfica. Mas ao contrário, existem caracteres anatômicos exclusivos de cada táxon, o que garante a eficácia dos mesmos para subsidiar decisões taxonômicas no grupo. v ABSTRACT The genus Ichnanthus P. Beauv. belongs to the tribe Paspaleae, subfamily Panicoideae (Poaceae), and currently includes ca. 33 species occurring predominantly in forests, especially in the Neotropical region. The genus has been traditionally recognized by leaf blades with asymmetrical base, paniculate inflorescences, spikelets laterally compressed, and the presence of winged appendages, scars or convex protuberances at the base of the upper lemma. One species of this genus, I. pallens (Sw.) Munro ex Benth., which possess scars at the base of the upper lemma, includes two varieties (I. pallens var. pallens and I. pallens var. major (Nees) Stieber). This species displays high variability in morphological vegetative and reproductive characters, with numerous morphotypes whose morphology often overlap between them and even with the morphology of related species such as I. tenuis (J.S Presl) Hitchc. and I. ruprechtii Döll; these three taxa constitute the I. pallens complex. Based on field-collected samples and material housed in herbaria we conduced morphoanatomical analyzes of leaves and inflorescences from taxa in this species complex, in order to find intra- and/or interspecific variations which could be useful to taxonomic decisions. For cross-sections of the leaf blades and inflorescence peduncles the samples were embedded in histological resin and analyzed using an optical microscope. For the study of leaf epidermis and anthecia surfaces we used both light microscopy and scanning electron microscopy. Leaf blades and inflorescences were also analyzed using a stereoscopic microscope. We observed considerable differences in the leaf anatomy of I. pallens when compared with the two other species analyzed, and three main patterns involving this species were observed: the „I. pallens sensu stricto’ pattern, comprising I. pallens var. major and I. pallens var. pallens modes axillaris, delicate and sensu stricto; the „I. pallens mode confertus‟, comprising exclusively I. pallens var. pallens mode vi confertus, and the „I. pallens mode areolatus, comprising I. var. pallens mode areolatus only. The other two species analyzed (I. tenuis and I. ruprechtii) also displayed distinct patterns. The information about the anatomy of the peduncle confirm these patterns, whereas macromorphological characters of the inflorescences and micromorphological characters of the anthecium surface often overlap. The variation found in morphoanatomical characters evaluated
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