Effects of nutrient addition and acidification on plant species diversity and seed germination in heathland.
The atmospheric deposition of sulfur and nitrogen compounds in the Netherlands has been responsible for decreasing plant species diversity in heathland. To unravel the relative importance of nitrogen compounds on soil acidification and eutrophication, and hence on the vegetation, we carried out a factorial addition experiment and a germination experiment in heathland on nutrient-poor sandy soil. We changed nutrient availability and acidity independently in eight different treatments that, respectively, added nutrients or carbon in various combinations (N, P, glucose) or added acidifying or neutralizing compounds. One treatment also involved adding Al. Additions occurred five times per year during 5 years, in an area from which sods had been removed before the experiment began. The same design was used for the germination experiment, but the treatments were applied for 2 years. Our results showed that acidification was the most important factor in reducing species diversity. In addition, the germination of several heathland species was significantly reduced in plots with a pH below 5, and germination was very poor in plots where Al had been added. The number of plant species declined particularly with increasing Al in the upper soil horizons. We conclude that this relationship is responsible for the influence of acidification on plant species richness in heathland. The influence of nutrient availability on species composition in heathland was subsidiary to acidity, but nutrient availability influenced species composition in an independent way. The growth of the three dominant species (Molinia caerulea, Calluna vulgaris and Erica tetralix) was limited by different nutrients. E. tetralix was limited by N, C. vulgaris by P and M. caerulea by both N and P. We argue that increased N availability will change the relative availability of N and P, which can decrease species diversity. Together these results show how factorial experiments can elucidate the complex ecological effects arising from sulfur and nitrogen deposition, revealing different mechanisms that change species richness and community composition.