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No Bioma Caatinga./ Daniela Santos Carneiro

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No Bioma Caatinga./ Daniela Santos Carneiro DANIELA SANTOS CARNEIRO TORRES DIVERSIDADE DE CROTON L. (EUPHORBIACEAE) NO BIOMA CAATINGA FEIRA DE SANTANA – BA 2009 UNIVERSIDADE ESTADUAL DE FEIRA DE SANTANA DEPARTAMENTO DE CIÊNCIAS BIOLÓGICAS PROGRAMA DE PÓS-GRADUAÇÃO EM BOTÂNICA DIVERSIDADE DE CROTON L. (EUPHORBIACEAE) NO BIOMA CAATINGA DANIELA SANTOS CARNEIRO TORRES Tese apresentada ao Programa de Pós- Graduação em Botânica da Universidade Estadual de Feira de Santana como parte dos requisitos para a obtenção do título de Doutora em Botânica. ORIENTADOR: PROFA. DRA. ANA MARIA GIULIETTI (UEFS) CO-ORIENTADORA: DRA. INÊS CORDEIRO (IBt) FEIRA DE SANTANA - BA 2009 Ficha Catalográfica – Biblioteca Central Julieta Carteado Torres, Daniela Santos Carneiro. T644d Diversidade de Croton (Euphorbiaceae) no bioma caatinga./ Daniela Santos Carneiro. – Feira de Santana, 2009. 296f.: il. tab., graf. Orientador: Ana Maria Giulietti Inês Cordeiro Tese (Doutorado em Botânica) – Universidade Estadual de Feira de Santana, 2009. BANCA EXAMINADORA _____________________________________________ Prof. Dr. Ricardo de Souza Secco _____________________________________________ Profa. Dra. Margareth Sales _____________________________________________ Prof. Dr. Luciano Paganucci de Queiroz _____________________________________________ Prof. Dr. Fávio França _____________________________________________ Prof(a). Dr(a). ANA MARIA GIULIETTI HARLEY Orientadora e Presidente da Banca Feira de Santana – BA 2009 Dedico aos meus filhos Breno e Guilherme, razão da minha vida, e ao meu marido Júnior, meu eterno amor. Aos meus pais, Antonio e Cleuza e meus irmãos Magaly, Herbert, Tatiana e Gerson, pelo amor, união e colaboração. “Não deixe de fazer algo que goste devido à falta de tempo, a única falta que terá será desse tempo que infelizmente não voltará mais”. (trecho do poema Vida de Mário Quintana) AGRADECIMENTOS No período de execução desse projeto encontrei muitas pessoas que direta ou indiretamente ajudaram na realização dessa obra. Assim agradeço. Primeiramente, agradeço à Universidade Estadual do Sudoeste da Bahia pela concessão da licença e do auxílio da bolsa de estudo fundamentais para que este trabalho fosse realizado. E os colegas da área de Botânica da UESB que cobriram a minha ausência nesse período. Ao Programa de Pós-Graduação em Botânica da Universidade Estadual de Feira de Santana, em especial ao Luciano Paganucci e a Adriana pelo suporte e pelo auxílio na participação de congressos e em algumas viagens de campo. Aos curadores dos herbários, pelo empréstimo das coleções e pela cordialidade com que fui recebida em suas instituições. À Fundação de Amparo à Pesquisa do Estado da Bahia (FAPESB), pelo auxílio financeiro para o desenvolvimento inicial desse projeto, principalmente para as coletas no Nordeste. Às minhas maravilhosas orientadoras, Ana Maria Giulietti e Inês Cordeiro, pela paciência, amizade, cumplicidade, confiança, orientação e incansável estímulo ao meu crescimento profissional e pessoal. Aos “crotonólogos” Ricarda Riina pelo apoio, correção e tradução do artigo das espécies novas, ao Paul Berry e Benjamin van Ee pela colaboração a este projeto, a Beatriz Caruzo pelo apoio, troca de experiências e auxílio nas coletas. À Luciano Paganucci, o meu eterno agradecimento por ter me socorrido no momento que mais precisei, muito obrigada pela maravilhosa colaboração, auxílio e encantamento pelos Croton. À Flávio França, meu eterno orientador, por ter me encaminhado ao estudo das Euphorbiaceae, à Alessandro Rapini pelo apoio, e por sempre estarem prontos a auxiliar. À Maria Regina pela hospitalidade e pelo auxílio nas coletas na Paraíba. À todos que pacientemente me acompanharam em expedições de coletas: Milene, Domingos, Rubens, Fábio, Elaine, Elnatan, Luciano, Flávio, Efigênia e Joanilson Júnior. Aos amigos com quem tive o prazer de conviver, alguns por mais tempo e com maior intensidade que outros, com os quais dividimos momentos de crescimento: Cosme, Élvia, Jomar, Silvia Karla, Déa, Ciça, Sil Ferreira, Ana Luiza, Fabrício, Domingos, Francisco Haroldo, Teo, Elaine, Marlon, Marla, Paty, Nara, Adriana, Léa e Luísa. Sei que sempre que precisar posso contar com vocês. À Ricardo (RickVB) pela disponibilidade e paciência em me socorrer para desvendar os mistérios do programa ArcView. Em especial, a Milene, amiga desde o início da minha caminhada botânica até os sofrimentos no concurso da UESB e nas disciplinas. Porém também compartilhamos muitas alegrias e angústias, e a Maria José (Zezé), minha mãezinha loira, pela amizade, conselhos e trocas de experiências que colaboraram para o meu equilíbrio. Sei que sem vocês seria ainda mais difícil terminar esse projeto. A toda a minha família, em especial aos meus pais e meus irmãos, pois sem o apoio e amor de vocês nada disso seria possível. Ofereço à vocês o meu eterno respeito e amor. Ao meu marido, Joanilson Júnior, por ter acreditado em mim, pelo amor, compreensão, admiração, paciência, consolos e conselhos, principalmente nos momentos mais difíceis e nas TPMs. À Breno, razão pela qual pude fazer esse doutorado, e a Gui que surgiu no finalzinho do caminho, vocês preenchem completamente a minha vida. Sem o apoio, incentivo e amor de vocês nada disso seria possível. Ofereço a vocês, esse trabalho com minha mais profunda gratidão. E finalmente agradeço a todos com o mais puro sentimento de alegria, amor e gratidão por ter concluído mais uma etapa da minha vida. Muitíssimo obrigada!!!!! Um cheiro. Caso tenha esquecido de alguém, é que minha memória não é mais a mesma, depois dessa experiência. RESUMO Croton L. possui ca. de 1.200 espécies pantropicais, com maior diversidade nas Américas. Esse estudo teve como objetivos o levantamento de Croton no bioma Caatinga, esclarecimento da circunscrição de suas espécies, a resolução de problemas taxonômicos, além de tentar elucidar a história biogeográfica e a sua diversificação na Caatinga. Este estudo foi baseado em coleções, observação das populações em campo, análise de coleções de herbário e análise filogenética associado a estimativa de tempo de divergência. Foram reconhecidas 68 espécies de Croton para a Caatinga, sendo 21 delas endêmicas desse bioma. Fazem parte deste trabalho: Chave de identificação, descrições, ilustrações, mapas de distribuição geográfica das espécies e até duas coleções de herbário, além de comentários sobre o tipo de vegetação e estado de conservação segundo os critérios da IUCN. Na taxonomia das espécies são indicados 16 lectótipos, um neótipo, oito sinônimos e descritas três novas espécies: Croton arenosus, C. glanduloso-bracteatus e C. harleyi. Também foi realizado um estudo filogenético baseado em dados moleculares (ITS e trnL-F) associado à estimativa de tempo de divergência. Nessa análise foram reconhecidas 22 linhagens de Croton na Caatinga, que teriam se originado há cerca de 23M.a.a., com diversificação entre 18-6M.a.a., no Mioceno. Essas linhagens, geralmente, estão associadas a espécies de outras áreas de florestas sazonalmente decíduas, sugerindo uma forte relação entre essas floras neotropicais. ABSTRACT Croton L. includes ca. 1,200 pantropical species with highest diversity in the Americas. The objectives of the current work are to survey the Croton species that occur in the Caatinga biome, to clarify the circumscription of the species, to solve taxonomic problems, besides to try to elucidate the biogeographic history of the genus and its diversification in the Caatinga. This study had as basis the visit to herbarium collections, observation of natural populations in the field, analyses of herbarium collections and phylogenetic analyses associated to estimations of divergence times. We identified 68 species of Croton that occur in the Caatinga, with 21 being endemic to the biome. This work comprises identification keys, descriptions, illustrations and maps of geographical distribution for the species, plus comments about the vegetation types and conservation status according to the IUCN. In the taxonomic treatment of the species we indicate 16 lectotypes, one neotype, 8 synonyms and we describe three new species: Croton arenosus, C. glanduloso-bracteatus and C. harleyi. We also conduced a phylogenetic study with basis on molecular data (ITS and trnL-F) associated to the estimative of divergence times. In this analysis we identified 22 different lineages of Croton in the Caatinga, whose origin is dated at about 23 m.y.a. and their diversification ocurred between 18 to 6 m.y.a. during the Miocene. These Caatinga lineages are usually associated to species from other areas of seasonally deciduous forestes, what suggests a strong relationship between these neotropical floras. SUMÁRIO AGRADECIMENTOS RESUMO I ABSTRACT II ÍNDICE DE TABELAS III ÍNDICE DE FIGURAS IV INTRODUÇÃO GERAL..........................................................................................22 Capítulo 1: Taxonomic notes in Croton (Euphorbiaceae) and three new species from the Brazilian Caatinga....................................................................................35 Capítulo 2: Diversidade de Croton L. (Euphorbiaceae) no Bioma Caatinga, Brasil......................................................................................................................59 Capítulo 3: Diversificação de Croton L. (Euphorbiaceae) e suas implicações para a biogeografia histórica do bioma Caatinga.........................................................198 CONCLUSÕES GERAIS.....................................................................................251 ANEXOS..............................................................................................................255
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