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Nas últimas duas décadas, estudos empíricos têm demonstrado a importância da facilitação entre plantas, especialmente sob condições ambientais severas. Tal constatação inspirou a proposição da hipótese do gradiente de estresse (HGE), um modelo conceitual que prediz que a freqüência relativa entre facilitação e competição deve variar inversarmente ao longo de um gradiente de estresse, sendo a facilitação mais comum sob condições extremamente severas. As restingas (ou planícies costeiras arenosas) são ambientes caracterizados por condições ambientais limitantes para o desenvolvimento das plantas, característica que as tornam um ambiente propício para o predomínio de facilitação. Além disso, o gradiente de intensidade de estresse e perturbação relacionado à distância do mar faz ainda deste ambiente um modelo ideal para testar a HGE.
O objetivo geral desta tese foi estudar facilitação entre plantas de restinga através dos seguintes objetivos específicos: i) Apresentar uma revisão sistemática dos trabalhos já publicados sobre facilitação entre plantas de restinga em escala mundial, assim como investigar, através de meta-análise, que fatores influenciam a presença e intensidade da facilitação entre plantas; ii) Testar se os padrões de associação espacial entre árvores adultas isoladas e plantas de distintas formas de vida se alteram ao longo de um gradiente ambiental praia-interior, conforme esperado pela HGE; iii) Testar a HGE através de experimentos de campo envolvendo transplante da espécie potencialmente beneficiada (espécie alvo), assim como manipulação de recurso limitante ao longo do gradiente praia-interior; iv) Testar se o sombreamento é um mecanismo de facilitação na restinga. Para os testes empíricos (objetivos ii-iv), as observações e experimentos foram realizados em um gradiente de restinga arbustiva do Parque Estadual da Ilha do Cardoso (PEIC) localizado em Cananéia, São Paulo, Brasil.
vizinho foi predominantemente negativa sobre o estágio de semente, mas positiva sobre a sobrevivência das plântulas; iv) O efeito de planta-berçário entre árvores da restinga é, pelo menos parcialmente, explicado pelo sombreamento gerado pela copa da espécie facilitadora.
Dessa forma, conclui-se que a facilitação via sombreamento é uma interação relevante entre árvores adultas e jovens na restinga. Entretanto, ao contrário do previsto pela HGE, o efeito de planta-berçário tende a ser mais intenso em condições ambientais mais amenas. Estes resultados sugerem a existência de um ciclo de retroalimentação positiva gerado a partir do estabelecimento das primeiras árvores isoladas. Este ciclo positivo tem implicações importantes para a compreensão da organização estrutural da vegetação da restinga arbustiva, em que moitas de espécies lenhosas estão distribuídas em uma matriz predominantemente herbácea. Adicionalmente, a facilitação entre árvores e sua dependência em relação às condições ambientais locais devem ser mecanismos essenciais para explicar o gradiente de fisionomias vegetacionais sobre os cordões arenosos, a qual inicia-se em uma vegetação aberta e termina em uma floresta, assim como para entender a dinâmica temporal desta vegetação ao longo do tempo. Por fim, os resultados também trazem implicações práticas, relacionadas ao uso potencial de plantas-berçário e sombra artificial como técnicas de manejo para aumentar o estabelecimento de mudas transplantadas em projetos de restauração de restingas arbustivo-arbóreas.
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In the last two decades, empirical studies have shown the importance of plant interaction, especially under harsh environmental conditions. These observations led to the proposition of the stress gradient hypothesis (SGH), a conceptual model predicting that the relative frequency of facilitation and competition will vary inversely across gradients of stress, with facilitation being the dominant interaction under highly stressful conditions. Coastal dunes (known as restinga in Brazil) are environments with limiting conditions for plant performance, which makes them suitable for facilitation predominance among plant interactions. Furthermore, there is a gradient of stress and disturbance related to the presence of the sea, making this environment an ideal model to test the SGH.
The general goal of this thesis was to study plant facilitation in coastal dunes with the following specific objectives: i) to perform a systematic review of published studies on plant facilitation in costal dunes around the world as well as to investigate, using a meta-analysis approach, factors that affect the existence and magnitude of plant facilitation; ii) to test if spatial association between isolated adult trees and understory plants of distinct life forms changes along a beach-to-inland gradient, as expected by the SGH; iii) to test the SGH through field experiments involving target species transplantation coupled with manipulation of limiting resource along a beach-to-inland gradient; iv) to test if shading is a facilitation mechanism in coastal dunes. The empirical tests (objectives ii-iv) were conducted in an open scrub coastal dune located at Ilha do Cardoso State Park, Cananéia, São Paulo, Brazil.
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