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UNIVERSIDADE ESTADUAL DE CAMPINAS Instituto De Biologia UNIVERSIDADE ESTADUAL DE CAMPINAS Instituto de Biologia TIAGO PEREIRA RIBEIRO DA GLORIA COMO A VARIAÇÃO NO NÚMERO CROMOSSÔMICO PODE INDICAR RELAÇÕES EVOLUTIVAS ENTRE A CAATINGA, O CERRADO E A MATA ATLÂNTICA? CAMPINAS 2020 TIAGO PEREIRA RIBEIRO DA GLORIA COMO A VARIAÇÃO NO NÚMERO CROMOSSÔMICO PODE INDICAR RELAÇÕES EVOLUTIVAS ENTRE A CAATINGA, O CERRADO E A MATA ATLÂNTICA? Dissertação apresentada ao Instituto de Biologia da Universidade Estadual de Campinas como parte dos requisitos exigidos para a obtenção do título de Mestre em Biologia Vegetal. Orientador: Prof. Dr. Fernando Roberto Martins ESTE ARQUIVO DIGITAL CORRESPONDE À VERSÃO FINAL DA DISSERTAÇÃO/TESE DEFENDIDA PELO ALUNO TIAGO PEREIRA RIBEIRO DA GLORIA E ORIENTADA PELO PROF. DR. FERNANDO ROBERTO MARTINS. CAMPINAS 2020 Ficha catalográfica Universidade Estadual de Campinas Biblioteca do Instituto de Biologia Mara Janaina de Oliveira - CRB 8/6972 Gloria, Tiago Pereira Ribeiro da, 1988- G514c GloComo a variação no número cromossômico pode indicar relações evolutivas entre a Caatinga, o Cerrado e a Mata Atlântica? / Tiago Pereira Ribeiro da Gloria. – Campinas, SP : [s.n.], 2020. GloOrientador: Fernando Roberto Martins. GloDissertação (mestrado) – Universidade Estadual de Campinas, Instituto de Biologia. Glo1. Evolução. 2. Florestas secas. 3. Florestas tropicais. 4. Poliploide. 5. Ploidia. I. Martins, Fernando Roberto, 1949-. II. Universidade Estadual de Campinas. Instituto de Biologia. III. Título. Informações para Biblioteca Digital Título em outro idioma: How can chromosome number variation indicate evolutionary relationships between the Caatinga, the Cerrado and the Atlantic forest? Palavras-chave em inglês: Evolution Dry forests Tropical forests Polyploidy Ploidy Área de concentração: Biologia Vegetal Titulação: Mestre em Biologia Vegetal Banca examinadora: Fernando Roberto Martins [Orientador] Ana Paula de Moraes Neiva Izabel Pierozzi Data de defesa: 27-02-2020 Programa de Pós-Graduação: Biologia Vegetal Identificação e informações acadêmicas do(a) aluno(a) - ORCID do autor: https://orcid.org/0000-0002-0244-9640 - Currículo Lattes do autor: http://lattes.cnpq.br/4756064884574811 Campinas, 27 e Fevereiro de 2020 COMISSÃO EXAMINADORA Prof. Dr. Fernando Roberto Martins Profa. Dra. Ana Paula de Moraes Profa. Dra. Neiva Izabel Pierozzi Os membros da Comissão Examinadora acima assinaram a Ata de Defesa, que se encontra no processo de vida acadêmica do aluno. A Ata da defesa com as respectivas assinaturas dos membros encontra-se no SIGA/Sistema de Fluxo de Dissertação/Tese e na Secretaria do Programa de Pós- graduação em Biologia Vegetal do Instituto de Biologia. AGRADECIMENTOS Ao meu orientador, Prof. Dr. Fernando Roberto Martins, pela paciência e empenho dedicado a minha formação e a este trabalho. Sinto-me privilegiado por ter tido mais um excelente orientador nesta jornada. À Profa. Dra. Eliana Regina Forni-Martins, pela contribuição, dedicação e paciência ao longo de todo o desenvolvimento deste projeto. Sou muito grato por todo conhecimento compartilhado. Aos professores membros da banca de qualificação, Prof. Dr. Louis Bernard Klaczko, Prof. Dr. Ricardo Augusto Lombello e Profa. Dra. Maria Fernanda Calió pelo paciente trabalho dе revisão e pelos questionamentos que aprimoraram deste trabalho. Aos professores membros da pré-banca, Dr. André Olmos Simões, Dra. Ana Paula de Moraes, Dra. Neiva Isabel Pierozzi pela revisão e sugestões. Aos membros da banca de defesa, Dra. Ana Paula de Moraes, Dra. Neiva Isabel Pierozzi, e aos suplentes, Dr. Ricardo Augusto Lombello e Dra. Luciana Bolsoni Lourenço Morandini, pela avaliação deste trabalho. Agradeço а todos оs professores dos Programas de Pós-Graduação em Biologia Vegetal e Ecologia pelо conhecimento compartilhado, pela dedicação а docência e pоr terem mе ensinado e mе feito aprender. Aos amigos que conheci nessa jornada, Cinthia Aparecida da Silva, Everton Alves Maciel, Laura Riba Hernández e Vinícius Londe, pela companhia durante minhas idas a Campinas. Um agradecimento especial à Cinthia por permitir que eu passasse as noites em sua casa quando necessário e pelo Vinícius por sempre tirar minhas dúvidas. A ambos pelos inúmeros conselhos. Aos inúmeros pesquisadores, brasileiros e estrangeiros, por compartilharem seu conhecimento nas redes sociais, permitindo que programas desafiadores como R e QGIS fossem mais fáceis de usar. Ao meu companheiro, Alexandre Ramos Rodrigues, que acompanha minha jornada desde a graduação. Pelo apoio incondicional, amor, carinho e paciência a todo o momento. Aos colegas do grupo de Whatsapp da BV, que sempre responderam as minhas perguntas. À Rebeca Politano Romanini, em especial, por me enviar referências. A todos citotaxonomistas, pois este trabalho não teria sido possível sem seus esforços em descrever e identificar os cromossomos das inúmeras espécies vegetais nos biomas estudados e no mundo. Aos funcionários da secretaria do Instituto de Biologia, pela paciência e profissionalismo. À minha família, por terem me dado condições para que me desenvolvesse academicamente. O presente trabalho foi realizado com apoio da Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Código de Financiamento 001. “Para onde vão os trens meu pai? Para Mahal, Tamí, para Camirí, espaços no mapa, e depois o pai ria: também pra lugar algum meu filho, tu pode ir e ainda que se mova o trem tu não te moves de ti.” Hilda Hilst RESUMO Neste trabalho testamos as hipóteses de que 1) a alta frequência de poliploides em um bioma poderia indicar que esse bioma tem origem recente e 2) poliploides são mais bem adaptados do que diploides a ambientes severos, sendo, portanto, mais frequentes nesses ambientes. Usamos a Caatinga, o Cerrado e a Mata Atlântica como biomas-modelo para testarmos estas hipóteses. Comparamos o número cromossômico das espécies presentes em cada bioma e a origem de alterações no número cromossômico (i.e., ocorrência de disploidia/aneuploidia e poliploidia) entre e dentro desses biomas. Esperamos que 1) se números cromossômicos mais altos e poliploides forem mais frequentes em ambientes severos, a Caatinga terá os maiores números de cromossomos e a maior proporção de poliploides, seguidos pelo Cerrado e pela Mata Atlântica; no entanto, 2) se números cromossômicos mais altos e poliploides forem mais frequentes em biomas mais recentes, o Cerrado terá os maiores números de cromossomos e a maior proporção de poliploides, seguidos pela Caatinga e pela Mata Atlântica. A partir de um extenso banco de dados florístico de espécies dicotiledôneas arbustivo- arbóreas que ocorrem em cada bioma, pesquisamos na literatura científica o número somático (2n) das espécies e o número base (x) de gêneros e famílias. Realizamos análises de distribuição de frequências (normalidade, assimetria e curtose) dos números cromossômicos em cada bioma, em espécies comuns aos três biomas e em espécies exclusivas de cada bioma; análise de variância (One- Way ANOVA) para comparar o número cromossômico médio entre cada bioma e entre espécies comuns e exclusivas; e análise de qui-quadrado de homogeneidade para comparar classes de número cromossômico entre biomas e entre espécies comuns e exclusivas e avaliar diferenças de origem das alterações do número cromossômico. Nosso banco de dados florístico contém 4092 espécies de dicotiledôneas arbustivo-arbóreas pertencentes a 826 gêneros e 126 famílias. Encontramos o número cromossômico de 965 espécies (23,6%) pertencentes a 412 gêneros e 82 famílias em publicações especializadas. Nossos dados revelaram que a distribuição da frequência dos números de cromossomos em todos os biomas e em espécies comuns e exclusivas era não-normal, leptocúrtica e moderadamente assimétrica para a direita. Entretanto, classes mais altas de números cromossômicos e poliploidia foram mais frequentes no Cerrado. Nossos resultados corroboram a hipótese de que a poliploidia pode indicar a origem mais recente de uma bioma. Palavras-chave: evolução, floresta seca, floresta úmida, ploidia, savana, disploidia. ABSTRACT It has been hypothesized that 1) high polyploidy frequency in a biome could indicate its recent origin and 2) that polyploids are better adapted to harsh environments than diploids, therefore being more frequent in such environments. In this work we tested for those hypotheses using the Atlantic forest, the Caatinga and the Cerrado as model biomes. We compared the chromosome number of species in each biome and the origin of chromosome number alteration (i.e, the occurrence of dysploidy/aneuploidy and polyploidy) among and within those biomes. We expected that 1) if high chromosome numbers and polyploidy are more frequent in severe environments, the Caatinga will have the highest chromosome numbers and the highest proportion of polyploids, followed by the Cerrado and the Atlantic forest; however, 2) if high chromosome numbers and polyploidy are more frequent in more recent biomes, the Cerrado will have the highest chromosome numbers and the highest proportion of polyploids, followed by the Caatinga and the Atlantic forest. From a floristic database of dicotyledonous shrub-tree species that occur in each biome, we surveyed the scientific literature for the somatic number (2n) of species and base number (x) of genera and families. We conducted analyses of frequency distribution (normality, skewness and kurtosis) of the chromosome numbers in each biome, of species common to all biomes and of species exclusive to each biome; analysis of variance (One- Way ANOVA), to compare the mean chromosome
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