“…'Nonforest species' are generally light-demanding species, and their richness in younger forest plots is likely to be due to the incomplete closure of the canopy, inertia of the understorey plant assemblage after recent canopy closure (grassland populations have been demonstrated to persist in forest for at least 50 years; Heubes et al, 2011), and to the persistence of altered soil properties due to the previous utilization as agricultural or grazing land (Matlack, 2009;Sciama et al, Figure 4 Results of variation partitioning for true forest species richness (TFS) in terms of fractions of variation explained by three groups of explanatory variables: forest successional age (> 77, > 56, > 38, > 22, < 22 years), environment/landscape variables (aspect, elevation, slope, mean annual temperature, mean maximum summer temperature, patch area, perimeterarea ratio, connectivity) and spatial variables (x, y, x 2 , y 2 , xy 2 and x 3 ). 'Nonforest species' are generally light-demanding species, and their richness in younger forest plots is likely to be due to the incomplete closure of the canopy, inertia of the understorey plant assemblage after recent canopy closure (grassland populations have been demonstrated to persist in forest for at least 50 years; Heubes et al, 2011), and to the persistence of altered soil properties due to the previous utilization as agricultural or grazing land (Matlack, 2009;Sciama et al, Figure 4 Results of variation partitioning for true forest species richness (TFS) in terms of fractions of variation explained by three groups of explanatory variables: forest successional age (> 77, > 56, > 38, > 22, < 22 years), environment/landscape variables (aspect, elevation, slope, mean annual temperature, mean maximum summer temperature, patch area, perimeterarea ratio, connectivity) and spatial variables (x, y, x 2 , y 2 , xy 2 and x 3 ).…”