Soil, biomass, and management of semi-natural vegetation. II. Factors controlling species diversity

Using a wide range of conditions and plant community types, species diversity was investigated in relation to edaphic and non-edaphic site conditions, management, and biomass characteristics. Both standing biomass and aboveground production were investigated, and their effects compared. Three taxonomic assemblages were studied: (1) vascular plants only, (2) bryophytes also included, (3) terrestrial lichens included as well. Using a multivariate approach, both species richness and evenness could best be explained when only vascular plants were considered, emphasizing the role of taxonomic restrictions. The models best explaining species richness merely required abiotic conditions. This supports recent theories emphasizing the importance of an environmental regulation of the pool of (adapted) species from which the actual species are recruited. Explanatory soil properties were moisture and pH (both unimodal), and the soil available N:P ratio. Plots with large perimeter:area ratios had significantly more species than those with low ratios, indicating the importance of consistency in quadrat shape. Hump-shaped species richness relationships could be identified for both standing biomass and productivity, but they explained only a small part of the variation and were apparent only if soil and management effects were not accounted for. Unimodality (and notably the decreasing phase) was most pronounced when using maximum standing biomass, suggesting that the key factor is competition for light. At intermediate levels of standing biomass, the positive effects of habitat productivity and the negative effects of standing biomass itself are in balance, and high species numbers may be expected. When soil or management variables were allowed in the models, hump-shaped biomass relationships were no longer confirmed, suggesting that such relationships may arise from the covariation of biomass with other factors. Management explained a much larger part of the variation than the hump-shaped biomass relationship, suggesting that mowing and hay removal (both showing independent positive effects) regulate species richness in ways other than solely through the control of maximum standing biomass. Significantly higher evenness values were obtained at sites with low maximum biomass values, but only for the vascular plants. In addition, species evenness was positively related to the frequency of mowing. The numbers of both rare and endangered species were strongly curtailed by high standing biomass values, suggesting that these species are more susceptible to competitive exclusion than others. Through direct as well as indirect effects, management is confirmed to be beneficial not only for general species richness, but also for the occurrence of rare and endangered species.