Diversity pattern and spatial scale: a study of a tropical rain forest of Malaysia

Scale is emerging as one of the critical problems in ecology because our perception of most ecological variables and processes depends upon the scale at which the variables are measured. A conclusion obtained at one scale may not be valid at another scale without sufficient knowledge of the scaling effect, which is also a source of misinterpretation for many ecological problems, such as the design of reserves in conservation biology. This paper attempts to study empirically how scaling may affect the spatial patterns of diversity (tree density, richness and Shannon diversity) that we may perceive in tropical forests, using as a test-case a 50 ha forest plot in Malaysia. The effect of scale on measurements of diversity patterns, the occurrence of rare species, the fractal dimension of diversity patterns, the spatial structure and the nearest-neighbour autocorrelation of diversity are addressed. The response of a variable to scale depends on the way it is measured and the way it is distributed in space. We conclude that, in general, the effect of scaling on measures of biological diversity is non-linear; heterogeneity increases with the size of the sampling units, and fine-scale information is lost at a broad scale. Our results should lead to a better understanding of how ecological variables and processes change over scale.