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Universidade Estadual De Campinas Instituto De Biologia UNIVERSIDADE ESTADUAL DE CAMPINAS INSTITUTO DE BIOLOGIA LUDMILA MARIA RATTIS TEIXEIRA ON THE HABITAT AND LANDSCAPES: THE EFFECT OF SCALE AND DATA ON SPECIES RESPONSE SOBRE O HÁBITAT E A PAISAGEM: EFEITOS DA ESCALA E DOS DADOS SOBRE A RESPOSTA DAS ESPÉCIES CAMPINAS 2016 LUDMILA MARIA RATTIS TEIXEIRA ON THE HABITAT AND LANDSCAPES: THE EFFECT OF SCALE AND DATA ON SPECIES RESPONSE SOBRE O HÁBITAT E A PAISAGEM: EFEITOS DA ESCALA E DOS DADOS SOBRE A RESPOSTA DAS ESPÉCIES Thesis presented to the Institute of Biology of the University of Campinas in partial fulfillment of the requirements for the degree of Doctor in Ecology Tese apresentada ao Instituto de Biologia da Universidade Estadual de Campinas como parte dos requisitos exigidos para a obtenção do Título de Doutora em Ecologia. ESTE ARQUIVO DIGITAL CORRESPONDE À VERSÃO FINAL DA TESE DEFENDIDA PELA LUDMILA MARIA RATTIS TEIXEIRA E ORIENTADA PELO PROF. DR. RAFAEL DIAS LOYOLA. Orientador: RAFAEL DIAS LOYOLA CAMPINAS 2016 Campinas, 16 de junho de 2016 COMISSÃO EXAMINADORA Prof. Dr. Rafael Dias Loyola Prof. Dr. Carlos Eduardo Viveiro Grelle Prof. Dr Paulo De Marco Junior Profa. Dra Eleonore Zulnara Freire Setz Prof. Dr. Leonardo Ré Jorge Os membros da Comissão Examinadora acima assinaram a Ata de defesa, que se encontra no processo de vida acadêmica do aluno. À Danielle, Nubia, Vanessa e Genevieve e a tudo que elas me ensinaram... Bate forte até sangrar a mão os tambores de Minas soarão seus tambores nunca se calarão... (Márcio Borges, mineiro) AGRADECIMENTOS Agradeço... ao Ensino Público Brasileiro, do qual desfrutei durante todo o ensino fundamental, parte do ensino médio e toda a Pós-Graduação; Ao Grupo Tal Pai Tal Filho, à Coordenação de Aperfeiçoamento Profissional do Ensino Superior e à Fundação de Amparo à Pesquisa do Estado de São Paulo pelo apoio financeiro; À Universidade Estadual de Campinas (UNICAMP), Universidade Federal de Goiás (UFG), Universidade Federal da Bahia (UFBA) e Carleton University, todos seus professores e funcionários; Ao Rafael Loyola por confiar em mim. O Rafa sabe que a Academia é formada por seres humanos, com problemas e idiossincrasias, e respeita cada um de seus colegas. Ele quem me encorajou a fazer os cursos de Frugivoria e Dispersão de Sementes e Análises Espaciais em Macroecologia, que foram tão importantes na minha formação como ecóloga. À Lenore Fahrig que me ensinou a remar, cortar lenha, acampar, acender fogueira, fazer bolo de chocolate na casca de laranja, assar marshmallows, testar hipóteses e escrever artigos científicos. No dia que entramos no barco para ela me ensinar a remar no Grant Lake, ela disse: eu direciono e você dá a força, nosso objetivo é alcançar aquela pedra, na outra margem... Obedeci e fizemos dois capítulos juntas durante o ano que estive na Carleton University. Aos membros da pré-banca: Sidney F. Gouveia, Marcus Cianciaruso e Divino Silvério pelas discussões e comentários. Aos membros da Banca Examinadora por aceitarem o convite e contribuírem para o crescimento deste trabalho. Ao Pedro Jordano por sua competência, elegância e simplicidade a ponto de “fazer ciência e soltar pipa” o que me faz lembrar outra pessoa a quem tenho muito a agradecer: meu avô científico Thomas Lewinsohn, o qual sempre me confundiu... Nunca soube dividir em quais momentos ele estava soltando pipa ou fazendo ciência, tamanha sua satisfação em lecionar Ecologia de Comunidades e discutir ciência nas horas que nunca são vagas quando se está ao seu lado. Ao Mauro Galetti pelas boas ideias durante o curso de Frugivoria e Dispersão de Sementes e às demais discussões que se seguiram. Ao Sérgio Furtado dos Reis pela compreensão e cordialidade. Ao Paulo de Marco que, no início de 2012, quando cheguei a Goiânia, ao mesmo tempo que o belo Otávio veio ao mundo, discutiu comigo meu trabalho, disse-me que, para ele, aluno de doutorado deveria ser tratado como um colega e, quando resolvi mudar-me para Goiânia me deu um forte abraço acolhedor. Valeu Paulo! À Sandra Martins, Ivan Correto, Poliana Mendes, Benjamin e Genevieve Perkins por terem cuidado de mim após meu acidente no Canadá. À Genevieve de novo, de novo e de novo. A Gen me ensinou a falar inglês, estudar estatística, tomar café e comer Vegemite, o que significa que devo tudo a ela. Ao Mafalda (Luis Francisco Mello Coelho) que sem dúvida é o responsável por minha entrada na Academia. Ao Leonardo Ré Jorge e Janaina Cortinóz por discutirem ciência e vida e me ajudarem nos trabalhos das disciplinas na Unicamp. Aos funcionários e professores de Carleton U e aos companheiros do Geomatics Landscape Ecology Lab pelas discussões e ajudas, em especial à Margareth, Fernanda Zimmerman, Sandra Martins, Sara Collins, Igor Pfeifer e Koreen Millard. Aos companheiros de UFG, do Laboratório de Biogeografia da Conservação (CB-Lab), Laboratório de Comunidades, LETS e The MetaLand, em especial à Fernanda Brum, Raísa Vieira, Lívia Laureto, Fernanda Fava, Leandro Maracahipes, Fernando Sobral, Marcos Carlucci, Ricardo Dobrovolski, Sidney Gouveia, Fabricio Villalobos, Diogo Samia, Luciano Sgarbi, Bruno Vilela, Miguel Ollala-Tárraga, Kelly Souza, Marcelo Weber, Welma, Daniel Paiva e Cristina Araújo. Cada uma dessas pessoas sabe o quanto me ajudou nesses quatro anos e agradeço a cada um por fazer parte da tese. À Capoeira Angola e a tudo que ela me deu pelas mãos dos Mestres Leninho e Guaxini do Mar. Ao pessoal do IPAM, em especial ao Paulo Brando pelas discussões, comentários e grande incentivo na reta final. Aos amigos com quem tive o prazer de dividir o grande teto dos alunos do programa de Ecologia da UFG, o edifício Barbosa Lima, com pessoas incríveis. Dentre eles, Tia Ísis, Eduardo Pacífico, Caroline Nóbrega, Sidney Gouveia, Fabricio Villalobos, Ricardo Dobrovolski, Diogo Provete, Bruno Vilela, Fábio Carvalho, Vanessa Staggemeier, Alexander Florez, Leandro Maracahipes, Thiago Bernardi, Poliana Mendes, Daniel Paiva e Karina Silva. Tenho que agradecer especialmente à Be e à Beubinha. Compartilhamos a essência, somos diferentes em todo o resto. Obrigada por dividirem comigo o café, vinho, o dramin e a co-autoria da filosofia “apamerrda”. Adoro discutir ciência com vocês! É sempre muito frutífero! Aos meus brilhantes amigos Gabriela Venturini e Vinicius Cardoso, pelo exemplo. À Danielle Miranda por discutir comigo o que realmente importa desde os tempos da graduação em Minas Gerais. À minha família, por ser tudo na minha vida. Eu sobrevivi e devo cada superação às orações dos meus avós Luzia e Oswaldo, à memória da vovó Geralda, à culinária do Rogerinho e à ajuda dos cunhados. Ao apoio incondicional do Paulinho Teixeira e da Fatinha Rattis, de seus filhos lindos e às filhas de seus filhos... Obrigada aos Teixeira por trabalharem tanto. Obrigada aos Rattis por acreditarem no impossível. ABSTRACT The environment of a given species is always changing and studying the causes and consequences of these changes is a challenging task in Ecology. This is because the very definition of suitable habitat is species-specific and even local-specific. Besides that, some factors can affect the results and conclusions coming from the studies carried out in landscapes. The habitat surrogate, the measurement of its effect on biodiversity, and the spatial extent of the investigation could change according to data availability and the purpose of research. Here we tested some hypothesis about the effect of habitat amount, patch size and isolation on species richness; spatial extent and species traits affecting the relationship between landscape structure and biodiversity and proposed a method to evaluate carrying capacity and connectivity on species’ range scale: (1) is habitat amount in the surrounding landscape as strong a predictor of species richness, as the combined predictive strength of patch isolation and patch size? (Chapter 1); (2) how does the scale of effect change with species traits and the measured population response? (Chapter 2); we also proposed a method to evaluate connectivity and carrying capacity at species range scale (Chapter 3). In chapter 1, our results supported the habitat amount hypothesis. Patch size and isolation did not have effects on species richness independent of habitat amount; they can be replaced by habitat amount alone. In chapter 2, we found that the spatial extent at which landscape structure mostly affect plant community depends on the movement range of the species, reproduction rate and the particular response analyzed. Species that disperse farther and produce more seeds showed a larger scale of effect, and species abundance responded to the surrounding landscape at a smaller scale than species occurrence. In Chapter 3, we highlighted that habitat amount and carrying capacity were more dependent on species’ features, whereas fragmentation and functional connectivity were more scale dependent, except when the species has high dispersal ability. The proposed approach can lead to lower risks of incurring in commission errors arising from landscape-scale underestimation of species’ occurrences and provide a new approach to assess species-habitat relationship. We also highlighted the importance of considering range-scaled landscape, landscape carrying capacity, and patch functional connectivity in a synthetic and integrated framework for conservation studies. Keywords: carrying capacity; deductive habitat suitability modeling; functional connectivity; habitat amount; herbaceous plant species; landscape ecology; scale of effect. RESUMO O ambiente das espécies está em constante mudança. Estudar as causas e consequências dessa mudança é uma das tarefas mais desafiadoras em Ecologia. Isto porque a definição de hábitat adequado é espécie-específica, podendo ser diferente até mesmo para uma mesma espécie que ocorre em diferentes locais. Além disso, alguns fatores podem afetar os resultados e conclusões adquiridas por meio de estudos sobre a mudança de hábitat na escala da paisagem. O conceito de hábitat, a avaliação do seu efeito sobre a biodiversidade e a extensão espacial do estudo podem mudar segundo a disponibilidade de dados e os objetivos almejados.
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