Species diversity of plant communities on calcareous screes: the role of intermediate disturbance

cover and on cover of individual species were obtained from 0.25 m 2 , 0.5 m 2 , and 1.0 m 2 plots. An index [100 – vegetation cover (%)] was used as a surrogate for direct quantification of disturbance (movement of stones and destruction of biomass). Its suitability was substantiated by the observation of position changes of marked stones after two years. In agreement with the intermediate disturbance hypothesis, significant unimodal responses of both species richness and species diversity to disturbance were found using plots of all three sizes. However, the maxima of species richness- and species diversity-disturbance curves were shifted towards higher vegetation cover (= lower disturbance) with increasing mean diameters of the scree-building stones. This shift can be interpreted either as a synergistic effect of disturbance frequency combined with disturbance intensity/extent, or as a result of an increase in spatial heterogeneity promoting species coexistence in successionally more advanced patches of vegetation. K e y w o r d s : Alps, patch dynamics, species coexistence, species richness, substrate instability, succession, successional mosaic