Roland Bobbink

Global assessment of nitrogen deposition effects on terrestrial plant diversity: a synthesis

Atmospheric nitrogen (N) deposition is a recognized threat to plant diversity in temperate and northern parts of Europe and North America. This paper assesses evidence from field experiments for N deposition effects and thresholds for terrestrial plant diversity protection across a latitudinal range of main categories of ecosystems, from arctic and boreal systems to tropical forests. Current thinking on the mechanisms of N deposition effects on plant diversity, the global distribution of G200 ecoregions, and current and future (2030) estimates of atmospheric N-deposition rates are then used to identify the risks to plant diversity in all major ecosystem types now and in the future. This synthesis paper clearly shows that N accumulation is the main driver of changes to species composition across the whole range of different ecosystem types by driving the competitive interactions that lead to composition change and/or making conditions unfavorable for some species. Other effects such as direct toxicity of nitrogen gases and aerosols, long-term negative effects of increased ammonium and ammonia availability, soil-mediated effects of acidification, and secondary stress and disturbance are more ecosystem- and site-specific and often play a supporting role. N deposition effects in mediterranean ecosystems have now been identified, leading to a first estimate of an effect threshold. Importantly, ecosystems thought of as not N limited, such as tropical and subtropical systems, may be more vulnerable in the regeneration phase, in situations where heterogeneity in N availability is reduced by atmospheric N deposition, on sandy soils, or in montane areas. Critical loads are effect thresholds for N deposition, and the critical load concept has helped European governments make progress toward reducing N loads on sensitive ecosystems. More needs to be done in Europe and North America, especially for the more sensitive ecosystem types, including several ecosystems of high conservation importance. The results of this assessment show that the vulnerable regions outside Europe and North America which have not received enough attention are ecoregions in eastern and southern Asia (China, India), an important part of the mediterranean ecoregion (California, southern Europe), and in the coming decades several subtropical and tropical parts of Latin America and Africa. Reductions in plant diversity by increased atmospheric N deposition may be more widespread than first thought, and more targeted studies are required in low background areas, especially in the G200 ecoregions.

Effects of nitrogen deposition and empirical nitrogen critical loads for ecoregions of the United States

Human activity in the last century has led to a significant increase in nitrogen (N) emissions and atmospheric deposition. This N deposition has reached a level that has caused or is likely to cause alterations to the structure and function of many ecosystems across the United States. One approach for quantifying the deposition of pollution that would be harmful to ecosystems is the determination of critical loads. A critical load is defined as the input of a pollutant below which no detrimental ecological effects occur over the long-term according to present knowledge. The objectives of this project were to synthesize current research relating atmospheric N deposition to effects on terrestrial and freshwater ecosystems in the United States, and to estimate associated empirical N critical loads. The receptors considered included freshwater diatoms, mycorrhizal fungi, lichens, bryophytes, herbaceous plants, shrubs, and trees. Ecosystem impacts included: (1) biogeochemical responses and (2) individual species, population, and community responses. Biogeochemical responses included increased N mineralization and nitrification (and N availability for plant and microbial uptake), increased gaseous N losses (ammonia volatilization, nitric and nitrous oxide from nitrification and denitrification), and increased N leaching. Individual species, population, and community responses included increased tissue N, physiological and nutrient imbalances, increased growth, altered root : shoot ratios, increased susceptibility to secondary stresses, altered fire regime, shifts in competitive interactions and community composition, changes in species richness and other measures of biodiversity, and increases in invasive species.

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.

A comparative study on nutrient cycling in wet heathland ecosystems

SummaryThe concept of the relative nutrient requirement (Ln) that was introduced in the first paper of this series is used to analyse the effects of the dominant plant population on nutrient cycling and nutrient mineralization in wet heathland ecosystems. A distinction is made between the effect that the dominant plant species has on (1) the distribution of nutrients over the plant biomass and the soil compartment of the ecosystem and (2) the recirculation rate of nutrients. The first effect of the dominant plant species can be calculated on the basis of the δ/k ratio (which is the ratio of the relative mortality to the decomposition constant). The second effect can be analysed using the relative nutrient requirement (Ln). The mass loss and the changes in the amounts of N and P in decomposing above-ground and below-ground litter produced by Erica tetralix and Molinia caerulea were measured over three years. The rates of mass loss from both above-ground and below-ground litter of Molinia were higher than those from Erica litter. After an initial leaching phase, litter showed either a net release or a net immobilization of nitrogen or phosphorus that depended on the initial concentrations of these nutrients. At the same sites, mineralization of nitrogen and phosphorus were measured for two years both in communities dominated by Molinia and in communities dominated by Erica. There were no clear differences in the nitrogen mineralization, but in one of the two years, phosphate mineralization in the Molinia-community was significantly higher. On the basis of the theory that was developed, mineralization rates and ratios between amounts of nutrients in plant biomass and in the soil were calculated on the basis of parameters that were independently measured. There was a reasonable agreement between predicted and measured values in the Erica-communities. In the Molinia-communities there were large differences between calculated and measured values, which was explained by the observation that the soil organic matter in these ecosystems still predominantly consisted of Erica-remains.

Differential effects of ammonium and nitrate on three heathland species

Arnica montana and Cirsium dissectum, typical of species-rich heathlands and acidic grasslands, have declined rapidly in The Netherlands in recent years. Field surveys suggest that the decline is caused by soil acidification as a result of enhanced atmospheric N and S deposition. Therefore, the survival, growth and development of these species were studied in a water culture experiment, using nutrient solutions which differed both in mineral nitrogen form and in ammonium concentration. For comparison, the performance of a third, acid tolerant species, Calluna vulgaris, was studied. The results showed that both Arnica and Cirsium performed better using nitrate than when using ammonium as a sole nitrogen source, whereas ammonium toxicity became apparent when ammonium concentrations were raised above 100 µM. Ammonium toxicity was expressed by an increase in mortality of Arnica plants with increasing ammonium concentrations and by a reduction of biomass in Arnica and Cirsium. Furthermore, cation concentrations in both roots and shoots decreased when ammonium was supplied as a nitrogen source. In contrast, Calluna showed optimal development when using ammonium as a sole nitrogen source. In this species, only root biomass was negatively affected by high ammonium concentrations. The ecological implications of these preferences are discussed in relation to soil acidification.

Nitrogen critical loads for natural and semi-natural ecosystems: The empirical approach

One of the major threats to the structure and the functioning of natural and semi-natural ecosystems is the recent increase in air-borne nitrogen pollution (NHy and NOx). Ecological effects of increased N supply are reviewed with respect to changes in vegetation and fauna in terrestrial and aquatic natural and semi-natural ecosystems. Observed and validated changes using data of field surveys, experimental studies or, of dynamic ecosystem models (the ‘empirical approach’), are used as an indication for the impacts of N deposition. Based upon these data N critical loads are set with an indication of the reliability. Critical loads are given within a range per ecosystem, because of spatial differences in ecosystems. The following groups of ecosystems have been treated: softwater lakes, wetlands & bogs, species-rich grasslands, heathlands and forests. In this paper the effects of N deposition on softwater lakes have been discussed in detail and a summary of the N critical loads for all groups of ecosystems is presented. The nitrogen critical load for the most sensitive ecosystems (softwater lakes, ombrotrophic bogs) is between 5–10 kg N ha−1 yr−1, whereas a more average value for the range of studied ecosystems is 15–20 kg N ha−1 yr−1. Finally, major gaps in knowledge with respect to N critical loads are identified.

Atmospheric deposition and canopy exchange processes in heathland ecosystems

The aims of the present study were to determine canopy exchange processes and to quantify total atmospheric deposition of sulphur and nitrogen in heathland. The study was carried out in dry inland heath vegetation, dominated by Calluna vulgaris, in two nature reserves in the eastern part of the Netherlands. Atmospheric deposition was determined with throughfall-stemflow measurements, adapted for low vegetation. Throughflow measurements (sum of throughfall and stemflow) in artificial Calluna canopies showed co-deposition of SOx and NHy upon heathland vegetation. In the real Calluna canopy, a significant part of the deposited ammonia/ammonium was directly assimilated by the Calluna shoots, especially in wet periods. The concentrations of potassium, calcium and magnesium in throughflow, after passage through the Calluna canopy, increased significantly compared with bulk precipitation. The amount of cations lost from the canopy were in good agreement with the observed ammonium uptake by the Calluna. A field experiment demonstrated that losses of the above-mentioned cations can be doubled by application of ammonium sulphate. It was shown that interception deposition is an important component of the atmospheric deposition of sulphur and nitrogen upon Calluna heathland; bulk precipitation amounted to only c. 35-40% of total atmospheric input. Total atmospheric deposition of sulphur and nitrogen in the investigated heathlands was 1.5-2.1 (27-33 kg S ha(-1) yr(-1)) and 2.1-3.1 kmolc ha(-1) yr(-1) (30-45 kg N ha(-1) yr(-1)), respectively. It is concluded that the present atmospheric nitrogen deposition is a continuous threat for the existence of heathlands in Western Europe.

Restoration management of abandoned chalk grassland in the Netherlands

After abandonment of chalk grassland in the Netherlands, Brachypodium pinnatum had become very dominant and this resulted in a dramatic decrease of plant diversity. Restoration of these abandoned sites is important, because of its former high diversity with many nationally endangered species. To restore the chalk grassland vegetation, the impact of the introduction of different experimental cutting regimes was investigated during three years. The thick litter layer and the dominance of Brachypodium was strongly reduced after introduction of all cutting regimes. In all cutting treatments forb phytomass increased considerably, especially in the twice-a-year cutting vegetation, where the forb phytomass became as high as that of Brachypodium. Phanerogamic species diversity was stimulated in all cutting regimes. Especially the number of short-lived forbs. with a persistent seed bank, increased markedly in the twice-a-year cutting treatment. It is concluded that cutting the vegetation twice is adequate to reduce the negative effects of the thick litter layer and the abundant growth of Brachypodium. It creates an appropriate starting point for more detinite management, whether mowing or grazing. However, the restoration of a more complete species assortment strongly depends on the re-invasion possibilities of these species.

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.

How nitrate leaching from agricultural lands provokes phosphate eutrophication in groundwater fed wetlands: the sulphur bridge

Increased phosphorus availability may provoke serious eutrophication problems in wetlands. Strong evidence indicates that sulphate induced mobilization of phosphate (internal eutrophication) has been responsible for a strong decline of the biodiversity in wetlands during the last decades. It is currently underestimated, however, that the wide spread leaching of nitrate from agricultural lands can indirectly provoke strong internal phosphate eutrophication in wetlands, via its interference with sulphur and iron biogeochemistry in the subsoil. Nitrate can mobilize sulphate from geological pyrite deposits by the oxidation of FeSx in the aquifer, leading to a decrease of nitrate and an increase of groundwater sulphate concentrations. Furthermore nitrate immobilizes iron in the subsoil by oxidizing reduced (dissolved) iron. Increased sulphate concentrations may provoke strong phosphate eutrophication in wetlands fed directly or indirectly (via surface water) with groundwater as sulphate strongly interferes with iron phosphorus chemistry and stimulates anaerobic decomposition of organic matter. Management of wetlands should therefore be approached at a broader scale which includes the landscape-scale management of groundwater systems. Leaching of nitrate to the groundwater, for instance, should not only receive attention for its potential effects on drinking water quality but above all because of the resulting large scale mobilization of sulphate from geological pyrite deposits and the immobilization of ferrous iron.