Instability on steep slopes mediates tree species co-existence in a warm–temperate mixed forest
We investigated the factors determining the distribution and dynamics of tree species in a warm–temperate mature mixed forest of evergreen coniferous and broad-leaved tree species in a steep mountainous area for 13 years in southwest Japan, with particular focus on instability of the ground surface. Among various site conditions, landform unit was the principal factor determining the distribution of tree species, while moisture regime was the second-most important factor within the upper area. The amount of movement of sediment and litter on the ground surface in the lower area was much higher than movement within the upper area, indicating that the lower area was unstable due to mass movement caused by erosion. The effects of instability of the ground surface on mortality and recruitment varied across the dominant tree species. Symplocos prunifolia (SYMPLOCACEAE), which was distributed in the upper area, and Machilus japonica, which was distributed in the lower area, exhibited lower mortality and higher recruitment in the areas where they were mainly distributed. These results suggest that topographic niche differentiation caused habitat segregation for some species. However, for most species, such relationships were not consistently observed, and growth rates did not significantly differ between the upper and lower areas. This study, by using long-term data, demonstrates that variation in sensitivity to stability due to topography contributes to local species richness and co-existence.