Ecological Factors Driving Tree Diversity across Spatial Scales in Temperate Forests, Northeast China

Species diversity has been shown to be influenced by environmental factors, but the mechanism underlying their relationship remains unclear across spatial scales. Based on field investigation data collected from 3077 sample plots in temperate forest ecosystems, we compared tree species richness, evenness and dominance at 10 km x 10 km, 30 km x 30 km and 90 km x 90 km spatial scales. Then, we detected the scale dependence of changes in tree species composition on climate, topography and forest structure using variation partitioning and quantified their contribution to tree diversity with gradient-boosted models (GBMs) and fitted their relationships. The magnitude of tree richness, evenness and dominance significantly increased with spatial scale. Ecological factors jointly accounted for 24.3%, 26.5% and 38.5% of the variation in tree species composition at the three spatial scales, respectively. The annual mean temperature had a strong impact on tree richness, evenness and dominance and peaked at an intermediate scale. Tree evenness and dominance increased with the variation of temperature but had upper and lower limits. Tree richness obviously increased with annual precipitation on multiple scales and decreased with annual sunshine duration at large spatial scales. Tree richness, evenness and dominance obviously increased with the variation in elevation and diameter at breast height at large scales and small scales, respectively. Tree dominance decreased with tree height at a small scale. The dependence of tree diversity on ecological factors increased with spatial scales. Furthermore, different factors exert various controls on tree diversity at different spatial scales, representing a comprehensive mechanism regulating tree diversity.