Plant species richness in mountain forests of the Bavarian Alps

Abstract

Based on a stratified random sample of 93 vegetation plots (144 m²) from montane and subalpine climax forests in a representative section through the Bavarian Alps, spatial pattern and environmental correlates of species density of trees, vascular understorey and epigeic bryophytes were analysed. Detecting landscape scale patterns in beta- and gamma-diversity based on interpretation of rarefaction curves proved to be difficult in a sample that had been stratified by ecological criteria. In 144 m² plots tree species density (5 ± 2.0, max. 10) declined with elevation and increased with stand age (multiple R ² = 0.557). The latter effect can be attributed to the secular history of game management and browsing pressure, which has hindered the regeneration of species-rich tree stands since ca. 150 yr. Species density of the forest undergrowth reached remarkably high levels for vascular plants (42 ± 12.8, max. 69) and bryophytes (14 ± 6.0, max. 30) and strongly depended on cover of the respective layer in a unimodal pattern, suggesting to separate direct and indirect effects, mediated through the mass effect, in the subsequent construction of regression models. Multiple regression (R ² = 0.47) revealed that vascular species density is limited chiefly through low plant cover, which in turn decreases with tree cover, elevation and soil quality, and secondly by species pools that contain larger numbers of species requiring high pH and ample light. Cover and direct effects had roughly equal weight in controlling bryophyte species density (R ² = 0.57). Biomass depended on the proportion of conifers in the tree layer and on site quality, less fertile sites tending to have higher bryophyte cover. The increase of bryophyte species density with elevation was interpreted as an effect of a pool of largely boreal-subalpine species. The increase of species density with stand age suggests dispersal limitation and deserves further study.