Tree competition and species coexistence in a warm-temperate old-growth evergreen broad-leaved forest in Japan

Abstract

The growth dynamics and mode of competition between adult trees ≥ 5.0cm in diameter at breast height (DBH) of nine abundant treespeciesoccupying ca. 85% of the total basal area were investigated in a 4ha study plot (200 m × 200 m) of awarm-temperate old-growth evergreen broad-leaved forest in the Tatera ForestReserve of Tsushima Island, southwestern Japan. In the plot, adult trees ≥5.0 cm DBH co-occurred with 35 woody plant species (except forwoody vine species). The most dominant and largest species,Castanopsis cuspidata var. sieboldiiexhibited a bimodal DBH distribution; it was found in both the upper and lowervertical layers. Other tree species had unimodal DBH distributionscorrespondingmostly to the lower vertical layer. We developed a model for individual growthincorporating both intra- and interspecific competition and degree ofcompetitive asymmetry. One-sided interspecific competition was detected in 17cases out of the 66 possible combinations on the scale of the 4 hastudy plot. The direction of interspecific competition was generally one-sidedfrom layer-I species to layer-II and III ones. The effects of two-sidedcompetition were detected only in layer-II and III species. OnlyDistylium racemosum exhibited one-sided intraspecificcompetition. We also found 11 cases of positive interspecific relationships.Generally, competitive relationships prevailed over positive relationshipsbetween adult trees in this warm-temperate evergreen broad-leaved forest.Competition between adult trees ≥ 5.0 cm in DBH did not occurinthe same vertical layer, but occurred only between trees in different verticallayers. This suggests that competition between adult trees ≥ 5.0cm in DBH plays a key role in the variation in species coexistencebetween different vertical layers on the 4 ha scale of thewarm-temperate evergreen broad-leaved forests. Moreover, it was found bycomparing with three different forest types that interspecific competition ismore intense in warm-temperate forests than in cool-temperate or sub-borealforests. We conclude that, compared to cool-temperate or sub-boreal forests(which have little interspecific competition), warm-temperate forests supportmore complex interspecific relationships and species-specific habitatpreferences that result in higher species diversity.