Distinguishing between turnover and nestedness in the quantification of biotic homogenization

Compositional changes through local extinction and colonization are inherent to natural communities, but human activities are increasingly influencing the rate and nature of the species being lost and gained. Biotic homogenization refers to the process by which the compositional similarity of communities increases over time through a non-random reshuffling of species. Despite the extensive conceptual development of the homogenization framework, approaches to quantify patterns of homogenization are scarcely developed. Most studies have used classical dissimilarity indices that actually quantify two components of compositional variation: turnover and nestedness. Here we demonstrate that a method that partitions those two components reveals patterns of homogenization that are otherwise obscured using traditional techniques. The forest understorey vegetation of an unmanaged reserve was recorded in permanent plots in 1979 and 2009. In only thirty years, the local species richness significantly decreased and the variation in the species composition from site to site shifted towards a structure with reduced true species turnover and increased dissimilarity due to nestedness. A classic analysis masked those patterns. In summary, we illustrated the need to move beyond the simple quantification of homogenization using classical indices and advocate integration of the multitude of ways to quantify community similarity into the homogenization framework.