Spatial segregation facilitates the coexistence of tree species in temperate forests

Competition between plants has an important role during the natural succession of forest communities. Niche separation between plants can reduce such interspecific competition and enable multispecies plant to achieve coexistence, although this proposition has rarely been supported in experiments. Plant competition can be captured by spatial segregation of the competing species to avoid fierce direct conflicts for nutrients and light. We investigated a site of 400 m 1000 m in Beijing Pine Mountain National Nature Reserve that was established for protecting Chinese pine and some rare fungi. Six dominant tree species (Fraxinus chinensis Roxb., Syringa reticulata (Blume) H. Hara var. amurensis (Rupr.) J. S. Pringle, Quercus mongolica Fisch. ex Ledeb., Armeniaca sibirica (L.) Lam., Pinus tabuliformis Carrière, and Ulmus pumila L.) were individually marked. Metrics of spatial segregation, based on the theory of spatial point process, were calculated to detect spatial competition. The corresponding type (species)-specific probabilities and the p-values from a spatially implicit test revealed significant overall spatial segregation between the six tree species. We further used the cross-type L-function to check the spatial correlation between Chinese pine and the other tree species, and detected a significant spatial repulsion relationship with four other tree species. Our study shows that each of the six dominant tree species occupies a different subarea in the landscape to effectively reduce direct spatial competition. We thus argue that patchy distributions of different tree species could be common in late forest community succession, and the coexistence of plants could be maintained over a large spatial scale. Management intervention, such as thinning the densities of dominant tree species, could be used to foster species coexistence and ensure the productivity of commercial stands