Species Richness Pattern along Altitudinal Gradient in Central European Beech Forests

The unimodal species richness-altitude distribution pattern seems to be universal. To investigate the validity of this phenomenon in homogeneous substrate and vegetation conditions, we sampled beech-dominated forests in five volcanic mountain ranges in the Western Carpathians. European beech (Fagus sylvatica L.) formed monodominant closed-canopy stands at altitudes from 300 to 1,200 m. Along this gradient, the influence of beech on understory plant species richness was expected to be strong and uniform. The shape of the species richness-altitude relationship was analyzed for three datasets: herb layer, shrub layer, and both layers merged together. Contrary to prediction, the studied species richness-altitude relationship was inversely unimodal, with a minimum at intermediate altitudes. Quadratic regression models were statistically significant for all three datasets (P < 0.001) and the explained variability ranged from 12 % to 20 %. The possible explanation for the observed pattern is twofold. In the central part of the altitudinal gradient, low species richness is due to strong competition by monodominant beech with accumulation of leaf litter and uptake soil resources, mainly water. This influence is somewhat released towards the margins of the gradient. Secondly, the species pool from the neighbouring communities increases species richness only in the lower parts of the altitudinal gradient.