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Biodiversidade De Clorófitas Marinhas Bentônicas Do Litoral Do Estado De São Paulo

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Biodiversidade De Clorófitas Marinhas Bentônicas Do Litoral Do Estado De São Paulo ANA CAROLINA SARAIVA PENA COTO Biodiversidade de Clorófitas Marinhas Bentônicas do Litoral do Estado de São Paulo Dissertação apresentada ao Instituto de Botânica da Secretaria do Meio Ambiente, como parte dos requisitos exigidos para a obtenção do título de MESTRE em BIODIVERSIDADE VEGETAL E MEIO AMBIENTE, na Área de Concentração de Plantas Avasculares e Fungos em Análises Ambientais. SÃO PAULO 2007 ANA CAROLINA SARAIVA PENA COTO Biodiversidade de Clorófitas Marinhas Bentônicas do Litoral do Estado de São Paulo Dissertação apresentada ao Instituto de Botânica da Secretaria do Meio Ambiente, como parte dos requisitos exigidos para a obtenção do título de MESTRE em BIODIVERSIDADE VEGETAL E MEIO AMBIENTE, na Área de Concentração de Plantas Avasculares e Fungos em Análises Ambientais. ORIENTADORA: DRA. DICLÁ PUPO SANTOS À Maria Aparecida Saraiva e José Manuel Pena Coto, meus pais, à Manuela, minha irmã e ao Ian, pela paciência, amor e incentivo. AGRADECIMENTOS À Dra. Diclá Pupo Santos pela orientação, confiança e oportunidade de aprendizado. À coordenação do Programa de Pós-Graduação do Instituto de Botânica. À CAPES pelo auxílio financeiro. À Dra. Nair S. Yokoya pela ajuda nos experimentos de cultura. À Dra. Andréa Tucci pelos conselhos e auxílio. À Dra. Mutue T. Fujii, Dra. Silvia M. P. B. Guimarães e Dra. Célia Sant’Anna pelos conselhos e disponibilidade em esclarecer dúvidas. Ao José Domingos e ao Ian pela ajuda nas coletas. À Manuela e Pierangeli pala ajuda na herborização do material. Ao Klei pela ajuda na confecção das pranchas de desenhos. À minha grande amiga Aline com quem sorri e chorei nestes últimos seis anos, pelo carinho, pelas conversas e desabafos diários. Às amigas Fernanda Ramlov e Suzana Fernandes, pela força, amizade e incentivo. Aos amigos Diógina, Valéria, Renato e Júlio, pelas dúvidas esclarecidas e pelas conversas engraçadas e descontraídas. À todos da seção de Ficologia do Instituto de Botânica. A Neide, a Neuzete, a Elizete, ao Manuel, pelo carinho, amizade e preocupação. Aos meus pais, José Manuel e Maria Aparecida, pelo amor incondicional, incentivo, carinho, paciência, orgulho e exemplo de garra e determinação. À Deus e à minha família, a quem sempre serei grata. Ao Ian, pelo amor, carinho, paciência, compreensão e incentivo nos principais momentos da minha vida. SUMÁRIO 1. INTRODUÇÃO.................................................................................................................. 01 2. OBJETIVOS....................................................................................................................... 05 3. ÁREA DE ESTUDO.......................................................................................................... 06 3.1. Caracterização da Área de Estudo ............................................................................. 06 3.2. Estações de Coleta...................................................................................................... 09 4. MATERIAIS E MÉTODOS............................................................................................... 12 4.1. Estudos Taxonômicos................................................................................................. 12 4.2. Cultura em Laboratório................................................................................................ 13 4.3. Comparação da flora atual de algas verdes marinhas com estudos anteriores............ 13 5. RESULTADOS................................................................................................................... 15 5.1. Ordem Ulvales............................................................................................................. 16 5.2. Ordem Cladophorales.................................................................................................. 42 5.3. Ordem Bryopsidales.................................................................................................... 81 5.4. Ordem Dasycladales................................................................................................... 103 Figuras......................................................................................................................... 106 5.5. Comparação da Flora................................................................................................... 135 6. DISCUSSÃO....................................................................................................................... 144 7. CONCLUSÃO..................................................................................................................... 149 8. BIBLIOGRAFIA................................................................................................................. 150 9. RESUMO/ ABSTRACT...................................................................................................... 163 1 1. INTRODUÇÃO A divisão Chlorophyta é complexa em termos de riqueza de espécies. Apresenta também grande diversidade no nível de organização do talo e histórico de vida de seus representantes. É o grupo de algas que mais se assemelha às plantas superiores, por apresentar clorofilas a e b como principais pigmentos fotossintetizantes, armazenar amido dentro de plastídeos, possuir pigmentos acessórios, tais como xantofila, luteína, zeaxantina, violaxantina e neoxantina e apresentar tilacóides dos cloroplastos agrupados em lamelas (Hoek et al. 1997). As células flageladas apresentam dois a quatro flagelos similares em estrutura, mas podendo diferir no tamanho. Alguns gêneros podem apresentar células flageladas estefanocontes, onde vários flagelos estão localizados na porção anterior da célula, formando uma coroa. Os cloroplastos estão envoltos por uma membrana dupla, não havendo membrana adicional do retículo endoplasmático. Os pirenóides, quando presentes, estão embutidos dentro do cloroplasto rodeados por grãos de amido (Hoek et al. 1997). Com base na estrutura do aparato flagelar, divisão celular e envoltório das células flageladas, Mattox & Stewart (1984) adotaram cinco classes para a divisão Chlorophyta: Micromonadophyceae, Charophyceae, Ulvophyceae, Pleurastrophyceae e Chlorophyceae. Lee (1989), além da ultraestrutura do aparato flagelar e dos tipos de envoltórios das células, considerou enzimas utilizadas na degradação de glicolato e uréia para definir quatro classes em Chlorophyta: Micromonadophyceae, Charophyceae, Ulvophyceae e Chlorophyceae. Hoek et al. (1997), com base na ultraestrutura das células flageladas, mitose e citocinese, histórico de vida e morfologia do talo, consideraram a divisão Chlorophyta com onze classes, cuja repartição foi fundamentada na idéia de que um organismo flagelado unicelular primitivo deu origem a várias linhagens de organismos unicelulares flagelados com diferenças ultraestruturais. Estes organismos teriam evoluído paralelamente, resultando na ocorrência de um mesmo nível de organização do talo em diferentes classes. Neste sistema de classificação, as 2 algas verdes marinhas bentônicas foram inseridas em quatro classes: Ulvophyceae, Cladophorophyceae, Bryopsidophyceae e Dasycladophyceae. Graham & Wilcox (2000) definem, para as algas verdes, duas linhagens multicelulares. A primeira compreende a classe Charophyceae, cujos representantes possuem aparato flagelar do tipo unilateral, com estrutura multiestratificada. A segunda linhagem engloba as classes Ulvophyceae, Trebouxiophyceae e Chlorophyceae, que apresentam aparato flagelar do tipo cruciado. As duas linhagens tiveram origem a partir de um organismo flagelado unicelular comum, próximo a Prasinophyceae, resultando em organismos multicelulares com os mesmos tipos de organização do talo, mas filogeneticamente distantes. Assim, os autores adotaram cinco classes para as verdes: Ulvophyceae, Trebouxiophyceae, Chlorophyceae, Prasinophyceae e Charophyceae. Este é o sistema de classificação que foi adotado no presente trabalho, onde as clorófitas marinhas foram circunscritas em Ulvophyceae, sendo a maioria de seus representantes incluída nas ordens Ulvales, Cladophorales, Bryopsidales e Dasycladales (Wynne 2005). A sistemática de Chlorophyta e sua filogenia, porém, continuam a ser amplamente discutidas, agora com base tanto em características morfológicas quanto em estudos moleculares (Zechman et al. 1990, Olsen et al. 1994, Woolcott et al. 2000, Leliaert et al. 2003, O’Kelly et al. 2004 e Lam & Zechman 2006). Estudos incluindo a ocorrência de clorófitas marinhas no Brasil foram citados desde Raddi (1823), Martius (1828), Martius et al. (1833) e Zeller (1876). Para o estado de São Paulo, as primeiras citações são de Lüderwaldt (1919) e Taylor (1930). Mas foi a partir da década de 50 que deu início um estudo mais efetivo, tanto das clorófitas marinhas, quanto de rodófitas e feófitas, com os trabalhos de flora de Joly (1957, 1965) realizados na Baía de Santos e arredores e no Litoral Norte e regiões circunvizinhas do Estado. Estas duas publicações representaram, junto com inúmeras adições à flora, uma nova era de conhecimento de algas no Brasil (Oliveira Filho 1977, Bicudo et al. 1998). 3 A partir daí, inúmeros levantamentos florísticos foram realizados no país, uns englobando todos os grupos de algas marinhas, como Ferreira-Correia et al. (1977), para o Maranhão, Ferreira & Pinheiro (1966), para o Ceará, Oliveira Filho & Ugadim (1976), para o Atol das Rocas, Nunes (1998), para a Bahia, Széchy et al. (1987, 1989), para o Arquipélago de Fernando de Noronha, Pedrini et al. (1989), Nassar (1994), Nassar et al. (2001) e Yoneshigue-Braga (1970), Yoneshigue
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