Effects of sampling scale on patterns of habitat association in tropical trees

Questions Niche differentiation is a central explanation for the co-existence and distribution patterns of numerous tree species in tropical forests, but functional equivalence leading to neutral dynamics has been proposed as an alternative explanation. This niche vs neutral debate is fuelled by the highly variable results yielded by studies of the association between tree species distributions and environmental factors, where some studies find strong associations but others do not. Here, we ask how differences in sampling scale between studies contribute to this variation. Location Barro Colorado Island, Panama. Methods Using distribution maps of canopy-statured individuals, we evaluated patterns of habitat association in five tropical tree species on Barro Colorado Island across a wide range of sampling scales (from 50 to 1600ha). We investigated the scale-dependency of species clumping patterns (Ripley's K) and the association of species distributions with important environmental variables (forest age, topography and geological formation) using point pattern analyses. Results Clump size and clump density had high variances within and among spatial scales. Significant habitat associations were found in all five species, with the number of habitat associations generally increasing with the sampling scale. Ignoring dispersal constraints inflated the number of significant habitat associations. Conclusions We found that patterns of habitat association (and hence conclusions on the importance of niche vs neutral processes) are strongly affected by the choice of sampling scale and location. Explicit inclusion of the effect of spatial scale is critical for studies of habitat association and the main processes that structure communities of tropical trees.