Effects of nutrient addition and acidification on plant species diversity and seed germination in heathland vegetation

1. The atmospheric deposition of sulphur 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. 2. 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. 3. 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. 4. 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. 5. 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. Erica tetralix was limited by N, Calluna vulgaris by P and Molinia 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. 6. Together these results show how factorial experiments can elucidate the complex ecological effects arising from sulphur and nitrogen deposition, revealing different mechanisms that change species richness and community composition

Nutrient limitation and botanical diversity in wetlands: Can fertilisation raise species richness?

The 'resource balance hypothesis' proposes that the species richness of grassland vegetation is potentially highest when the N:P ratio of plant tissues is 10-15 (co-limitation), so that species richness could be raised by fertilisation with N or P at sites with lower or higher N:P ratios, respectively. Here we use data from field surveys in Swiss, Dutch and American fens or wet grasslands to analyse what changes in N:P ratios might produce noticeable changes in species richness. Plant species numbers, above-ground biomass, tissue N and P concentrations and soil pH were recorded in plots of 0.06-4 m2. In each data set, plots with intermediate tissue N:P ratios (6-20) were on average most species-rich, but N:P ratios explained only 5-37% of the variation in species richness. Moreover, these effects were partially confounded with those of vegetation biomass and/or soil pH. The unique effects of N:P ratios (excluding those shared with biomass and pH) explained 11-17% of variation in species richness. The relationship between species richness and N:P ratios was asymmetric: plots with high N:P ratios were more species-poor than those with low N:P ratios. This was paralleled by a smaller species pool size at high N:P ratios (estimated from species numbers in multiple records), suggesting that fewer species are adapted to P-limited conditions than to N-limited conditions. According to these data, species richness in wetlands may possibly be raised by P-fertilisation when the initial N:P ratio of the vegetation is well above 20, but this option is not recommended for nature conservation as it might promote common species at the expense of rare one