Plant community responses to experimental climate manipulation in a Welsh ombrotrophic peatland and their palaeoenvironmental context.

We test whether vegetation community composition from a 10-year climate manipulation experiment on a Welsh peat bog resembles vegetation communities during periods of climate change inferred from a peat core. Experimentally warmed and combined warmed and droughted treatments drove significant increases in ericaceous shrubs but Sphagnum was unaffected. Similarly, Calluna vulgaris seeds increase during inferred warmer periods in the palaeoecological record. Experimental short-term episodic drought (four 4-week drought treatments) did not affect vegetation. Plant community composition has undergone several abrupt changes throughout the past c. 1500 years, often in response to human disturbance. Only slight changes occurred during the Medieval Climate Anomaly (c. 950-1250 Common Era [CE]) in vegetation and hydrology, while abrupt changes occurred during the Little Ice Age (c. 1300-1850 CE) when water tables were highest, suggesting that these shifts were driven by changes in water table, modulated by climate. A period of water table drawdown c. 1800, synchronous with historical records of increased drainage, corresponds with the development of the present-day vegetation community. Modern analogues for fossil material, characterized by abundant Rhynchospora alba and Sphagnum pulchrum, are more common after this event. Vegetation changes due to climate inferred from the palaeo record differ from those observed in the experiments, possibly relating to differences in the importance of drivers of vegetation change over varying timescales. Whereas temperature is frequently identified as the dominant driver of plant community change in experiments, sustained changes in water table appear to be more important in the long-term record. We find evidence that recent climate change and other anthropogenic stressors (e.g. drainage, heavy metal and nitrogen pollution) may promote the development of novel plant communities without analogues in the fossil record. These communities may be poorer at sequestering carbon and may respond differently to future climate change.

Impact of nitrogen deposition at the species level.

In Europe and, increasingly, the rest of the world, the key policy tool for the control of air pollution is the critical load, a level of pollution below which there are no known significant harmful effects on the environment. Critical loads are used to map sensitive regions and habitats, permit individual polluting activities, and frame international negotiations on transboundary air pollution. Despite their fundamental importance in environmental science and policy, there has been no systematic attempt to verify a critical load with field survey data. Here, we use a large dataset of European grasslands along a gradient of nitrogen (N) deposition to show statistically significant declines in the abundance of species from the lowest level of N deposition at which it is possible to identify a change. Approximately 60% of species change points occur at or below the range of the currently established critical load. If this result is found more widely, the underlying principle of no harm in pollution policy may need to be modified to one of informed decisions on how much harm is acceptable. Our results highlight the importance of protecting currently unpolluted areas from new pollution sources, because we cannot rule out ecological impacts from even relatively small increases in reactive N deposition.

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.

A synthesis of methane emissions from 71 northern, temperate, and subtropical wetlands.

Wetlands are the largest natural source of atmospheric methane. Here, we assess controls on methane flux using a database of approximately 19 000 instantaneous measurements from 71 wetland sites located across subtropical, temperate, and northern high latitude regions. Our analyses confirm general controls on wetland methane emissions from soil temperature, water table, and vegetation, but also show that these relationships are modified depending on wetland type (bog, fen, or swamp), region (subarctic to temperate), and disturbance. Fen methane flux was more sensitive to vegetation and less sensitive to temperature than bog or swamp fluxes. The optimal water table for methane flux was consistently below the peat surface in bogs, close to the peat surface in poor fens, and above the peat surface in rich fens. However, the largest flux in bogs occurred when dry 30-day averaged antecedent conditions were followed by wet conditions, while in fens and swamps, the largest flux occurred when both 30-day averaged antecedent and current conditions were wet. Drained wetlands exhibited distinct characteristics, e.g. the absence of large flux following wet and warm conditions, suggesting that the same functional relationships between methane flux and environmental conditions cannot be used across pristine and disturbed wetlands. Together, our results suggest that water table and temperature are dominant controls on methane flux in pristine bogs and swamps, while other processes, such as vascular transport in pristine fens, have the potential to partially override the effect of these controls in other wetland types. Because wetland types vary in methane emissions and have distinct controls, these ecosystems need to be considered separately to yield reliable estimates of global wetland methane release.

Mapping Portuguese Natura 2000 sites in risk of biodiversity change caused by atmospheric nitrogen pollution.

In this paper, we assess and map the risk that atmospheric nitrogen (atN) pollution poses to biodiversity in Natura 2000 sites in mainland Portugal. We first review the ecological impacts of atN pollution on terrestrial ecosystems, focusing on the biodiversity of Natura 2000 sites. These nature protection sites, especially those located within the Mediterranean Basin, are under-characterized regarding the risk posed by atN pollution. We focus on ammonia (NH3) because this N form is mostly associated with agriculture, which co-occurs at or in the immediate vicinity of most areas of conservation interest in Portugal. We produce a risk map integrating NH3 emissions and the susceptibility of Natura 2000 sites to atN pollution, ranking habitat sensitivity to atN pollution using expert knowledge from a panel of Portuguese ecological and habitat experts. Peats, mires, bogs, and similar acidic and oligotrophic habitats within Natura 2000 sites (most located in the northern mountains) were assessed to have the highest relative risk of biodiversity change due to atN pollution, whereas Natura 2000 sites in the Atlantic and Mediterranean climate zone (coastal, tidal, and scrubland habitats) were deemed the least sensitive. Overall, results allowed us to rank all Natura 2000 sites in mainland Portugal in order of evaluated risk posed by atN pollution. The approach is of great relevance for stakeholders in different countries to help prioritize site protection and to define research priorities. This is especially relevant in countries with a lack of expertise to assess the impacts of nitrogen on biodiversity and can represent an important step up from current knowledge in such countries.

Author Correction: Nitrogen-rich organic soils under warm well-drained conditions are global nitrous oxide emission hotspots.

The original version of this Article contained an error in the first sentence of the Acknowledgements section, which incorrectly referred to the Estonian Research Council grant identifier as "PUTJD618". The correct version replaces the grant identifier with "PUTJD619". This has been corrected in both the PDF and HTML versions of the Article.

Nitrogen-rich organic soils under warm well-drained conditions are global nitrous oxide emission hotspots.

Nitrous oxide (N2O) is a powerful greenhouse gas and the main driver of stratospheric ozone depletion. Since soils are the largest source of N2O, predicting soil response to changes in climate or land use is central to understanding and managing N2O. Here we find that N2O flux can be predicted by models incorporating soil nitrate concentration (NO3-), water content and temperature using a global field survey of N2O emissions and potential driving factors across a wide range of organic soils. N2O emissions increase with NO3- and follow a bell-shaped distribution with water content. Combining the two functions explains 72% of N2O emission from all organic soils. Above 5 mg NO3--N kg-1, either draining wet soils or irrigating well-drained soils increases N2O emission by orders of magnitude. As soil temperature together with NO3- explains 69% of N2O emission, tropical wetlands should be a priority for N2O management.

Long-term nitrogen deposition increases heathland carbon sequestration.

The large increases in reactive nitrogen (N) deposition in developed countries since the Industrial Revolution have had a marked impact on ecosystem functioning, including declining species richness, shifts in species composition, and increased N leaching. A potential mitigation of these harmful effects is the action of N as a fertiliser, which, through increasing primary productivity (and subsequently, organic matter production), has the potential to increase ecosystem carbon (C) storage. Here we report the response of an upland heath to 10years of experimental N addition. We find large increases in plant and soil C and N pools, with N-driven C sequestration rates in the range of 13-138kgCkg-1. These rates are higher than those previously found in forest and lowland heath, mainly due to higher C sequestration in the litter layer. C sequestration is highest at lower N treatments (10, 20, and 40kgNha-1yr-1 above ambient), with evidence of saturation at the highest N treatment, reflecting a physiologically aged Calluna vulgaris (Calluna) canopy. To maintain these rates of sequestration, the Calluna canopy should be managed to maximise it's time in the building phase. Scaling our results across UK heathlands, this equates to an additional 0.77Mt CO2e per annum extra C sequestered into plant litter and the top 15cm of heathland soil as a result of N deposition. The bulk of this is found in the litter and organic soil horizons that hold an average of 23% and 54% of soil C, respectively. This additional C represents around 0.44% of UK annual anthropogenic GHG emissions. When considered in the context of falling biodiversity and altered species composition in heathland, policy focus should remain on reducing N emissions.

Taxonomic and functional turnover are decoupled in European peat bogs.

In peatland ecosystems, plant communities mediate a globally significant carbon store. The effects of global environmental change on plant assemblages are expected to be a factor in determining how ecosystem functions such as carbon uptake will respond. Using vegetation data from 56 Sphagnum-dominated peat bogs across Europe, we show that in these ecosystems plant species aggregate into two major clusters that are each defined by shared response to environmental conditions. Across environmental gradients, we find significant taxonomic turnover in both clusters. However, functional identity and functional redundancy of the community as a whole remain unchanged. This strongly suggests that in peat bogs, species turnover across environmental gradients is restricted to functionally similar species. Our results demonstrate that plant taxonomic and functional turnover are decoupled, which may allow these peat bogs to maintain ecosystem functioning when subject to future environmental change.

Evidence for differential effects of reduced and oxidised nitrogen deposition on vegetation independent of nitrogen load.

Nitrogen (N) deposition impacts natural and semi-natural ecosystems globally. The responses of vegetation to N deposition may, however, differ strongly between habitats and may be mediated by the form of N. Although much attention has been focused on the impact of total N deposition, the effects of reduced and oxidised N, independent of the total N deposition, have received less attention. In this paper, we present new analyses of national monitoring data in the UK to provide an extensive evaluation of whether there are differences in the effects of reduced and oxidised N deposition across eight habitat types (acid, calcareous and mesotrophic grasslands, upland and lowland heaths, bogs and mires, base-rich mires, woodlands). We analysed data from 6860 plots in the British Countryside Survey 2007 for effects of total N deposition and N form on species richness, Ellenberg N values and grass:forb ratio. Our results provide clear evidence that N deposition affects species richness in all habitats except base-rich mires, after factoring out correlated explanatory variables (climate and sulphur deposition). In addition, the form of N in deposition appears important for the biodiversity of grasslands and woodlands but not mires and heaths. Ellenberg N increased more in relation to NHx deposition than NOy deposition in all but one habitat type. Relationships between species richness and N form were habitat-specific: acid and mesotrophic grasslands appear more sensitive to NHx deposition while calcareous grasslands and woodlands appeared more responsive to NOy deposition. These relationships are likely driven by the preferences of the component plant species for oxidised or reduced forms of N, rather than by soil acidification.

Nitrogen deposition and reduction of terrestrial biodiversity: evidence from temperate grasslands.

Biodiversity is thought to be essential for ecosystem stability, function and long-term sustainability. Since nitrogen is the limiting nutrient for plant growth in many terrestrial ecosystems, reactive nitrogen has the potential to reduce the diversity of terrestrial vegetation and associated biota through favouring species adapted to quickly exploiting available nutrients. Although the potential has long been recognised, only recently has enough evidence come together to show beyond reasonable doubt that these changes are already occurring. Linked together, experimental, regional/empirical, and time-series research provide a powerful argument that enhanced deposition of reactive nitrogen across Great Britain, and potentially the rest of Europe, has resulted in a significant and ongoing decline in grassland species richness and diversity.

Consequences of human modification of the global nitrogen cycle.

The demand for more food is increasing fertilizer and land use, and the demand for more energy is increasing fossil fuel combustion, leading to enhanced losses of reactive nitrogen (Nr) to the environment. Many thresholds for human and ecosystem health have been exceeded owing to Nr pollution, including those for drinking water (nitrates), air quality (smog, particulate matter, ground-level ozone), freshwater eutrophication, biodiversity loss, stratospheric ozone depletion, climate change and coastal ecosystems (dead zones). Each of these environmental effects can be magnified by the 'nitrogen cascade': a single atom of Nr can trigger a cascade of negative environmental impacts in sequence. Here, we provide an overview of the impact of Nr on the environment and human health, including an assessment of the magnitude of different environmental problems, and the relative importance of Nr as a contributor to each problem. In some cases, Nr loss to the environment is the key driver of effects (e.g. terrestrial and coastal eutrophication, nitrous oxide emissions), whereas in some other situations nitrogen represents a key contributor exacerbating a wider problem (e.g. freshwater pollution, biodiversity loss). In this way, the central role of nitrogen can remain hidden, even though it actually underpins many trans-boundary pollution problems.

Impacts of atmospheric pollution on the plant communities of British acid grasslands.

Air pollutants are recognised as important agents of ecosystem change but few studies consider the effects of multiple pollutants and their interactions. Here we use ordination, constrained cluster analysis and indicator value analyses to identify potential environmental controls on species composition, ecological groupings and indicator species in a gradient study of UK acid grasslands. The community composition of these grasslands is related to climate, grazing, ozone exposure and nitrogen deposition, with evidence for an interaction between the ecological impacts of base cation and nitrogen deposition. Ozone is a key agent in species compositional change but is not associated with a reduction in species richness or diversity indices, showing the subtly different drivers on these two aspects of ecosystem degradation. Our results demonstrate the effects of multiple interacting pollutants, which may collectively have a greater impact than any individual agent.

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.

Regional trends in soil acidification and exchangeable metal concentrations in relation to acid deposition rates.

The deposition of high levels of reactive nitrogen (N) and sulphur (S), or the legacy of that deposition, remain among the world's most important environmental problems. Although regional impacts of acid deposition in aquatic ecosystems have been well documented, quantitative evidence of wide-scale impacts on terrestrial ecosystems is not common. In this study we analysed surface and subsoil chemistry of 68 acid grassland sites across the UK along a gradient of acid deposition, and statistically related the concentrations of exchangeable soil metals (1 M KCl extraction) to a range of potential drivers. The deposition of N, S or acid deposition was the primary correlate for 8 of 13 exchangeable metals measured in the topsoil and 5 of 14 exchangeable metals in the subsoil. In particular, exchangeable aluminium and lead both show increased levels above a soil pH threshold of about 4.5, strongly related to the deposition flux of acid compounds.

A classification and regression tree model of controls on dissolved inorganic nitrogen leaching from European forests.

Often, there is a non-linear relationship between atmospheric dissolved inorganic nitrogen (DIN) input and DIN leaching that is poorly captured by existing models. We present the first application of the non-parametric classification and regression tree approach to evaluate the key environmental drivers controlling DIN leaching from European forests. DIN leaching was classified as low (<3), medium (3-15) or high (>15kg N ha(-1) year(-1)) at 215 sites across Europe. The analysis identified throughfall NO(3)(-) deposition, acid deposition, hydrology, soil type, the carbon content of the soil, and the legacy of historic N deposition as the dominant drivers of DIN leaching for these forests. Ninety four percent of sites were successfully classified into the appropriate leaching category. This approach shows promise for understanding complex ecosystem responses to a wide range of anthropogenic stressors as well as an improved method for identifying risk and targeting pollution mitigation strategies in forest ecosystems.

Nitrogen deposition and reduction of terrestrial biodiversity: evidence from temperate grasslands.

Biodiversity is thought to be essential for ecosystem stability, function and long-term sustainability. Since nitrogen is the limiting nutrient for plant growth in many terrestrial ecosystems, reactive nitrogen has the potential to reduce the diversity of terrestrial vegetation and associated biota through favouring species adapted to quickly exploiting available nutrients. Although the potential has long been recognised, only recently has enough evidence come together to show beyond reasonable doubt that these changes are already occurring. Linked together, experimental, regional/empirical, and time-series research provide a powerful argument that enhanced deposition of reactive nitrogen across Great Britain, and potentially the rest of Europe, has resulted in a significant and ongoing decline in grassland species richness and diversity.