Individual species-area relationship of woody plant communities in a heterogeneous subtropical monsoon rainforest.

The spatial structure of species richness is often characterized by the species-area relationship (SAR). However, the SAR approach rarely considers the spatial variability of individual plants that arises from species interactions and species' habitat associations. Here, we explored how the interactions of individual plants of target species influence SAR patterns at a range of neighborhood distances. We analyzed the data of 113,988 woody plants of 110 species from the Fushan Forest Dynamics Plot (25 ha), northern Taiwan, which is a subtropical rainforest heavily influenced by typhoons. We classified 34 dominant species into 3 species types (i.e., accumulator, repeller, or no effect) by testing how the individual species-area relationship (i.e., statistics describing how neighborhood species richness changes around individuals) of target species departs (i.e., positively, negatively, or with no obvious trend) from a null model that accounts for habitat association. Deviation from the null model suggests that the net effect of species' interactions increases (accumulate) or decreases (repel) neighborhood species richness. We found that (i) accumulators were dominant at small interaction distances (<10-30 m); (ii) the detection of accumulator species was lower at large interaction distances (>30 m); (iii) repellers were rarely detected; and (iv) large-sized and abundant species tended to be accumulators. The findings suggest that positive species interactions have the potential to accumulate neighborhood species richness, particularly through size- and density-dependent mechanisms. We hypothesized that the frequently disturbed environment of this subtropical rainforest (e.g., typhoon-driven natural disturbances such as landslides, soil erosion, flooding, and windthrow) might create the spatial heterogeneity of species richness and promote positive species interactions.