Habitat specialization and phylogenetic structure of tree species in a coastal Brazilian white-sand forest

Aims The coastal Brazilian rainforest on white-sand(restinga)ranks among the most fragmented forest types in the tropics,owing to both the patchy distribution of sandy soils and widespread coastal development activities.Here we study the environmental and evolutionary determinants of a forest tree assemblage at a single restinga forest in Southeastern Brazil.We also explore the ability of competing hypotheses to explain the maintenance of species diversity in this forest type,which includes contrasting extremes of edaphic conditions associated with flooding stress.Methods The study was conducted in a white-sand forest permanent plot of 10.24 ha on the coastal plain of Southeastern Brazil.This plot was divided into 256 quadrats of 20×20 m,which were classified into two main edaphic habitats(flooded and drained).Trees with a diameter≥1 cm at breast height were identified.We assembled DNA sequence data for each of the 116 morphospecies recognized using two chloroplast markers(rbcL and matK).A phylogenetic tree was obtained using the maximum likelihood method,and a phylogenetic distance matrix was produced from an ultrametric tree.We analyzed similarity in floristic composition and structure between habitats and related them to cross-plot distances using permutation procedures.Null model torus shift simulations were performed to obtain a statistical significance level for habitat association for each species.The phylogenetic structure for the two habitats and for each 20×20 m quadrat was calculated using the mean phylogenetic distance weighted by species abundance and checked for significance using the standardized effect size generated by 5000 randomizations of phylogenetic tip labels.Important Findings Our results indicate that partitioning among edaphic habitats is important for explaining species distributions and coexistence in restinga forests.Species distributions within the plot were found to be non-random:there was greater floristic similarity within than between habitats,and>40%of the more abundant species were positively or negatively associated with at least one habitat.Patterns of habitat association were not independent of phylogenetic relatedness:the community was overdispersed with respect to space and habitat type.Closely related species tended to occur in different habitats,while neighboring trees tended to belong to more distantly related species.We conclude that habitat specialization is important for the coexistence of species in restinga forests and that habitat heterogeneity is therefore an essential factor in explaining the maintenance of diversity of this unique but fragile and threatened type of forest.