Variation of plant diversity in a temperate unmanaged forest in northern Spain: behind the environmental and spatial explanation

The persistence of seedlings in the forest understorey is of major importance for the maintenance and regeneration of canopy trees in several forested ecosystems. In the present study, we examine the small-scale spatial pattern of a mixed beech and oak seedling–sapling bank in two areas of an unmanaged temperate deciduous forest with different environmental conditions. We used environmental, biotic and spatial variables to establish the main factors that explain the spatial pattern of these seedling–sapling banks at different scales. The stand structure in both areas was similar, but while in plot A beech dominated the canopy, plot B was dominated by oaks. In both areas, established beech individuals showed a clear reverse J-shaped distribution, whereas established oaks showed a unimodal distribution with only a few young individuals. Seedlings of beech and oak were distributed in aggregates, whereas beech saplings had a random distribution. At broader scales, the abundance of seedlings and saplings is affected by the environment as well as by inter-species competition, while at finer scales the spatial pattern is mainly influenced by stochastic processes, probably related to seed predation and establishment. The structure of the seedling–sapling bank indicates an advantage of beech over oak as far as regeneration is concerned. Beech seedlings and saplings tolerate the stress induced by the canopy and the understorey and persist for many years, while oak seedlings decline in a few years. Therefore, if current conditions persist, after canopy opening beech seedlings and saplings can grow rapidly into the canopy and the stands will move towards beech dominance.