Atmospheric nitrogen deposition is related to plant biodiversity loss at multiple spatial scales.

Due to various human activities, including intensive agriculture, traffic, and the burning of fossil fuels, in many parts of the world, current levels of reactive nitrogen emissions strongly exceed pre-industrial levels. Previous studies have shown that the atmospheric deposition of these excess nitrogen compounds onto semi-natural terrestrial environments has negative consequences for plant diversity. However, these previous studies mostly investigated biodiversity loss at local spatial scales, that is, at the scales of plots of typically a few square meters. Whether increased atmospheric nitrogen deposition also affects plant diversity at larger spatial scales remains unknown. Here, using grassland plant community data collected in 765 plots, across 153 different sites and 9 countries in northwestern Europe, we investigate whether relationships between atmospheric nitrogen deposition and plant biodiversity are scale-dependent. We found that high levels of atmospheric nitrogen deposition were associated with low levels of plant species richness at the plot scale but also at the scale of sites and regions. The presence of 39% of plant species was negatively associated with increasing levels of nitrogen deposition at large (site) scales, while only 1.5% of the species became more common with increasing nitrogen deposition, indicating that large-scale biodiversity changes were mostly driven by "loser" species, while "winner" species profiting from high N deposition were rare. Some of the "loser" species whose site presence was negatively associated with atmospheric nitrogen deposition are listed as "threatened" in at least some EU member states, suggesting that nitrogen deposition may be a key contributor to their threat status. Hence, reductions in reactive nitrogen emissions will likely benefit plant diversity not only at local but also at larger spatial scales.

Soil phosphorus constrains biodiversity across European grasslands.

Nutrient pollution presents a serious threat to biodiversity conservation. In terrestrial ecosystems, the deleterious effects of nitrogen pollution are increasingly understood and several mitigating environmental policies have been developed. Compared to nitrogen, the effects of increased phosphorus have received far less attention, although some studies have indicated that phosphorus pollution may be detrimental for biodiversity as well. On the basis of a dataset covering 501 grassland plots throughout Europe, we demonstrate that, independent of the level of atmospheric nitrogen deposition and soil acidity, plant species richness was consistently negatively related to soil phosphorus. We also identified thresholds in soil phosphorus above which biodiversity appears to remain at a constant low level. Our results indicate that nutrient management policies biased toward reducing nitrogen pollution will fail to preserve biodiversity. As soil phosphorus is known to be extremely persistent and we found no evidence for a critical threshold below which no environmental harm is expected, we suggest that agro-environmental schemes should include grasslands that are permanently free from phosphorus fertilization.

Ecosystem responses to reduced and oxidised nitrogen inputs in European terrestrial habitats.

While it is well established that ecosystems display strong responses to elevated nitrogen deposition, the importance of the ratio between the dominant forms of deposited nitrogen (NH(x) and NO(y)) in determining ecosystem response is poorly understood. As large changes in the ratio of oxidised and reduced nitrogen inputs are occurring, this oversight requires attention. One reason for this knowledge gap is that plants experience a different NH(x):NO(y) ratio in soil to that seen in atmospheric deposits because atmospheric inputs are modified by soil transformations, mediated by soil pH. Consequently species of neutral and alkaline habitats are less likely to encounter high NH(4)(+) concentrations than species from acid soils. We suggest that the response of vascular plant species to changing ratios of NH(x):NO(y) deposits will be driven primarily by a combination of soil pH and nitrification rates. Testing this hypothesis requires a combination of experimental and survey work in a range of systems.

The impact of nitrogen deposition on acid grasslands in the Atlantic region of Europe.

A survey of 153 acid grasslands from the Atlantic biogeographic region of Europe indicates that chronic nitrogen deposition is changing plant species composition and soil and plant-tissue chemistry. Across the deposition gradient (2-44 kg N ha(-1) yr(-1)) grass richness as a proportion of total species richness increased whereas forb richness decreased. Soil C:N ratio increased, but soil extractable nitrate and ammonium concentrations did not show any relationship with nitrogen deposition. The above-ground tissue nitrogen contents of three plant species were examined: Agrostis capillaris (grass), Galium saxatile (forb) and Rhytidiadelphus squarrosus (bryophyte). The tissue nitrogen content of neither vascular plant species showed any relationship with nitrogen deposition, but there was a weak positive relationship between R. squarrosus nitrogen content and nitrogen deposition. None of the species showed strong relationships between above-ground tissue N:P or C:N and nitrogen deposition, indicating that they are not good indicators of deposition rate.

Nitrogen deposition threatens species richness of grasslands across Europe.

Evidence from an international survey in the Atlantic biogeographic region of Europe indicates that chronic nitrogen deposition is reducing plant species richness in acid grasslands. Across the deposition gradient in this region (2-44 kg N ha(-1) yr(-1)) species richness showed a curvilinear response, with greatest reductions in species richness when deposition increased from low levels. This has important implications for conservation policies, suggesting that to protect the most sensitive grasslands resources should be focussed where deposition is currently low. Soil pH is also an important driver of species richness indicating that the acidifying effect of nitrogen deposition may be contributing to species richness reductions. The results of this survey suggest that the impacts of nitrogen deposition can be observed over a large geographical range.

Decline of acid-sensitive plant species in heathland can be attributed to ammonium toxicity in combination with low pH.

The effects of increasing ammonium concentrations in combination with different pH levels were studied on five heathland plant species to determine whether their occurrence and decline could be attributed to ammonium toxicity and/or pH levels. Plants were grown in growth media amended with four different ammonium concentrations (10, 100, 500 and 1000 micromol l(-1)) and two pH levels resembling acidified (pH 3.5 or 4) and weakly buffered (pH 5 or 5.5) situations. Survival of Antennaria dioica and Succisa pratensis was reduced by low pH in combination with high ammonium concentrations. Biomass decreased with increased ammonium concentrations and decreasing pH levels. Internal pH of the plants decreased with increasing ammonium concentrations. Survival of Calluna vulgaris, Deschampsia flexuosa and Gentiana pneumonanthe was not affected by ammonium. Moreover, biomass increased with increasing ammonium concentrations. Biomass production of G. pneumonanthe reduced at low pH levels. A decline of acid-sensitive species in heathlands was attributed to ammonium toxicity effects in combination with a low pH.