Growth responses of low-alpine dwarf-shrub heath species to nitrogen deposition and management

Nitrogen deposition is a continuing problem in European alpine regions. We hypothesised that, despite climatic limitations, low-alpine Calluna heathland would respond to nitrogen addition with increased shoot growth and flowering and that fire and grazing would modify responses. In a five-year study, 0-50kgNha(-1)y(-1) were added, combined with burning (+/-) and clipping (+/-). Calluna vulgaris responded with increased shoot extension, but effects on flowering were variable. Burning enhanced the positive effect of nitrogen addition and negative effects of clipping. Sub-dominant shrubs generally did not respond to nitrogen. C. vulgaris shoot extension was stimulated by nitrogen addition of 10kgNha(-1)y(-1) (above background) supporting suggestions that alpine heathlands are sensitive to low levels of nitrogen deposition. Increased C. vulgaris growth could negatively impact on important lichen components of this vegetation through increased shading and competition. Climatic factors constrain productivity in this community, but do not prevent rapid responses to nitrogen deposition by some species.     

Sulphur and seasalt deposition as reflected by throughfall and runoff chemistry in forested catchments

At forested catchments at Lake G?rdsj?n on the Swedish west coast the deposition and runoff chemistry has been followed during the period 1979-1990 by throughfall and runoff measurements as well as by measurements of atmospheric concentrations. The 10-year means in throughfall and runoff are very similar for sulphur and the main seasalt ions sodium and chloride; for sulphur 26.1 and 27.6 kg ha(-1) yr(-1), for sodium 49 and 52 kg ha(-1) yr(-1) and for chloride 96 kg ha(-1) yr(-1) and 93 kg ha(-1) yr(-1), respectively. The actual flows are 100-200% higher than the wet deposition as collected in open bulk precipitation collectors indicating a very large input by dry deposition. One important question is to what extent the throughfall and runoff values can be used as measures of total deposition. We present results from studies at different experimental catchments illustrating the possibilities of using throughfall and runoff data as measures of atmospheric deposition of sulphur and seasalt.     

Deposition of fixed atmospheric nitrogen and foliar nitrogen content of bryophytes and Calluna vulgaris (L.) Hull

Atmospheric deposition of fixed nitrogen as nitrate and ammonium in rain and by dry deposition of nitrogen dioxide, nitric acid and ammonia has increased throughout Europe during the last two decades, from 2-6 kg N ha(-1) year(-1) to 15-60 kg N ha(-1) year(-1). The nitrogen contents of bryophytes and the ericaceous shrub Calluna vulgaris have been measured at a range of sites, with the objective of showing the degree to which nitrogen deposition is reflected in foliar plant nitrogen. Tissue nitrogen concentrations of herbarium bryophyte samples and current samples of the same species collected from the same sites were compared. No significant change in tissue nitrogen was recorded at a remote site in north-west Scotland where nitrogen inputs are small (< 6 kg N ha(-1) year(-1)). Significant increases in tissue N occurred at four sites ranging from 38% in central Scotland to 63% in Cumbria where nitrogen inputs range from 15 to 30 kg N ha(-1) year(-1). The relationships found between the estimated input of atmospheric nitrogen and the tissue nitrogen content of the selected bryophytes and Calluna at the sites investigated were found to be generally linear and fitted the form N(tissue) = 0.62 + 0.022 N(dep) for bryophytes and N(tissue) = 0.83 + 0.045 N(dep) for Calluna. There was thus an increase in total tissue nitrogen of 0.02 mg g(-1) dry weight for bryophytes and 0.045 mg g(-1) dry weight for Calluna for an increase in atmospheric nitrogen deposition of 1 kg ha(-1) year(-1). The lowest concentrations were found in north-west Scotland and the highest in Cumbria and the Breckland heaths of East Anglia, both areas of high atmospheric nitrogen deposition (30-40 kg N ha(-1) year(-1)). The implications of increased tissue nitrogen content in terms of vegetation change are discussed. Changes in atmospheric nitrogen deposition with time were also examined using measured values and values inferred from tissue nitrogen content of mosses. The rate of increase in nitrogen deposition is not linear over the 90-year period, and the increases were negligible over the period 1880-1915. However, during the period 1950 to 1990 the data suggest an increase in nitrogen deposition of 2 kg N ha(-1) every 10 years.     

Effect of vegetation type on throughfall deposition and seepage flux

This paper compares different vegetation types (coniferous and deciduous forest, grassed and pure heathland) in terms of input (throughfall deposition) and output (seepage flux) in a region with intermediate nitrogen load (+/-20kg Nha(-1)y(-1) via bulk precipitation) in comparable conditions in north Belgium. Coniferous forest (two plots Pinus sylvestris and two plots Pinus nigra) received significantly higher nitrogen and sulphur throughfall deposition than deciduous forest and heathland. Grassed and pure heathland had significantly highest throughfall quantities of Ca(2+) and Mg(2+), respectively. The observed differences in throughfall deposition between the different vegetation types were not univocally reflected in the ion seepage flux. Considerable seepage fluxes of NO(3)(-), SO(4)(2-), Ca(2+) and Al(III) were only found under the P. nigra plots. We discuss our hypothesis that the P. nigra forests already evolved to a situation of N saturation, while the other vegetation types did not.     

Short- and medium-term effects of experimental nitrogen fertilization on arthropods associated with Calluna vulgaris heathlands in north-west Spain

We studied the short- and medium-term effects of experimental nitrogen fertilization (3 and 15 months after the treatment) on the arthropods of Calluna vulgaris heathlands in NW Spain. Three heathland sites were selected with two permanent plots per site: control and fertilized. Ammonium nitrate fertilizer (56 kg N ha(-1) yr(-1)) was applied monthly and insects were caught using pitfall traps. We found mainly species-level responses to nitrogen addition. Seven species (e.g. Lochmaea suturalis) showed a consistent trend (benefited or harmed) in both periods and were proposed as possible reliable indicators of the effects of nitrogen deposition in these ecosystems. We also found variable arthropod trophic-group responses: (a) herbivores (leaf beetles, true bugs) increased in abundance on a short-term scale; (b) predators (carabid beetles, true bugs) showed opposite and less clear responses in both periods. Further long-term studies are needed to determine the mechanisms underlying the observed arthropod responses.     

Spatial variation in atmospheric nitrogen deposition on low canopy vegetation

Current knowledge about the spatial variation of atmospheric nitrogen deposition on a local scale is limited, especially for vegetation with a low canopy. We measured nitrogen deposition on artificial vegetation at variable distances of local nitrogen emitting sources in three nature reserves in the Netherlands, differing in the intensity of agricultural practices in the surroundings. In the nature reserve located in the most intensive agricultural region nitrogen deposition decreased with increasing distance to the local farms, until at a distance of 1500 m from the local nitrogen emitting sources the background level of 15 kg N ha(-1) yr(-1) was reached. No such trend was observed in the other two reserves. Interception was considerably lower than in woodlands and hence affected areas were larger. The results are discussed in relation to the prospects for the conservation or restoration of endangered vegetation types of nutrient-poor soil conditions.     

Reduced nitrogen has a greater effect than oxidised nitrogen on dry heathland vegetation

We investigated the effects of different ratios of reduced (NH4+) versus oxidised (NO3(-)) nitrogen in deposition on heathland and species-rich grassland vegetation at high nitrogen deposition levels in large mesocosms filled with nutrient-poor soils to which different NH4+/NO3(-) ratios were applied. The response of the forbs, Antennaria dioica, Arnica montana, Gentiana pneumonanthe, Thymus serpyllum, the grasses Danthonia decumbens, Deschampsia flexuosa, Nardus stricta and the shrub Calluna vulgaris was recorded. The forb A. dioica and the grass D.decumbens preferred low NH4+/NO3(-) ratios and were characterised by a negative correlation between NH4+/NO3(-) ratios and biomass and survival, whereas the grasses N. stricta and D. flexuosa showed no correlation with NH4+/NO3(-) ratios. Lime addition eliminated the negative effects of high NH4+ concentrations in deposition for A. dioica and the grass D. decumbens. The implications of these findings for heathland vegetations are discussed.     

Eutrophication trends in forest soils in Galicia (NW Spain) caused by the atmospheric deposition of nitrogen compounds

We calculated the sensitivity of Galician forest soils to eutrophication caused by atmospheric deposition of nitrogen compounds, using the Simple Mass Balance (SMB) method as described by [Posch, M., de Vries, W., Hettelingh, J.-P., 1995. Critical loads of sulphur and nitrogen. In: Posch, M., de Smet, P.A.M., Hettelingh, J.-P., Downing, R.J. Calculation and Mapping of Critical Thresholds in Europe. Status Report 1995, Coordination Center for Effects, National Institute for Public Health and the Environment, Bilthoven, The Netherlands, pp. 31-42]. Deposition values were used to calculate critical loads exceedance. Galician natural forest ecosystems can support nitrogen deposition loads of more than 10 kg Nha (-1) yr (-1). The lowest critical loads (approximately 10 kg Nha (-1) yr (-1)) mainly occurred in forest stands in the interior zone, while highest critical load values (approximately 68 kg Nha (-1) yr (-1)) were observed in eucalyptus stands at low altitudes in the littoral area. Exceedances based on N deposition levels, calculated from data recorded in 2001, occurred in 40% of the forest soils, showing the need to control N emissions in these areas to prevent possible eutrophication of soils and waters. Analysis of rainfall bulk composition revealed that ammonium, probably derived from agricultural and cattle activities, was the main compound responsible for N deposition in Galicia.     

Phosphorus supply influences heathland responses to atmospheric nitrogen deposition

On an upland moor dominated by pioneer Calluna vulgaris and with an understorey of mosses and lichens, experimental plots were treated with factorial combinations of nitrogen (N) at +0 and +20kg Nha(-1)yr(-1), and phosphorus (P) at +0 and +5kg Pha(-1)yr(-1). Over the 4-year duration of the experiment, the cover of the Calluna canopy increased in density over time as part of normal phenological development. Moss cover increased initially in response to N addition but then remained static; increases in cover in response to P addition became stronger over time, eventually causing reductions in the cover of the dominant Calluna canopy. Lichen cover virtually disappeared within 4 years in plots receiving +20kg Nha(-1)yr(-1) and also in separate plots receiving +10kg Nha(-1)yr(-1), but this effect was reversed by the addition of P.     

Dry deposition and internal circulation of nitrogen,sulphur and base cations to a coniferous forest

The dry deposition of sulphur, nitrogen and base cations to a spruce stand was estimated during a five year period using a surrogate surface resembling needles, throughfall and bulk deposition measurements. The deposition was calculated from the ratio between the deposition of an ion and sodium on the surrogate surface and the net throughfall of sodium to the forest. The dry deposition represented a large fraction of the total atmospheric input of base cations. For Na⁺, Mg²⁺, Ca²⁺, and K⁺ they were 66, 67, 53 and 42%, respectively. The internal circulation was 95% of non-marine net throughfall fro K⁺ and 76% for Ca²⁺. The dry deposition of SO₂ to the canopies regulates the internal circulation of Ca²⁺. The dry deposition of SO₂ to the canopies regulates the internal circulation of Ca²⁺. The dry depositions of ammonium and nitrate are close to the net throughfall of Kjeldahl-N and nitrate, respectively. The obtained deposition velocities are comparable to other studies. The calculated dry deposition of ammonium was compared to the net throughfall of ammonium at three nearby forest stands receiving different ammonium amounts on the soils. No correlation to nitrogen level was found, but most ammonium was lost and converted to organic nitrogen in the canopies of the wettest forest stand.     

Nitrogen deposition in Mediterranean forests

Atmospheric deposition of inorganic nitrogen was studied at two forested sites in the Montseny mountains (northeast Spain), peripheral to the Barcelona conurbation, and at a nearby lowland town, using bulk deposition, wet-only deposition, throughfall, and dry deposition inferred from branch-washes and surrogate surfaces (metacrylate plates). Bulk deposition inputs of ammonium and nitrate did not show significant temporal trends over a 16-year period. Bulk inputs of inorganic N were moderate, ranging from 6 to 10 kg N ha(-1) year(-1) depending on the time period considered and the degree of site exposure to polluted air masses from the Barcelona conurbation. Large dry-sedimented particles played a minor role, since wet-only inputs were virtually identical to bulk inputs. On the contrary, branch- and plate-washes indicated substantial dry inputs of N gases and small particles. Total atmospheric deposition was estimated at 15-22 kg N ha(-1) year(-1), most of it being retained within the studied broadleaved evergreen forests. Ecosystem N availability is thus likely to be increasing in these forests.     

Stress responses of Calluna vulgaris to reduced and oxidised N applied under 'real world conditions'

Effects and implications of reduced and oxidised N, applied under 'real world' conditions, since May 2002, are reported for Calluna growing on an ombrotrophic bog. Ammonia has been released from a 10 m line source generating monthly concentrations of 180-6 microg m(-3), while ammonium chloride and sodium nitrate are applied in rainwater at nitrate and ammonium concentrations below 4mM and providing up to 56 kg N ha(-1) year(-1) above a background deposition of 10 kg N ha(-1) year(-1). Ammonia concentrations, >8 microg m(-3) have significantly enhanced foliar N concentrations, increased sensitivity to drought, frost and winter desiccation, spring frost damage and increased the incidence of pathogen outbreaks. The mature Calluna bushes nearest the NH3 source have turned bleached and moribund. By comparison the Calluna receiving reduced and oxidised N in rain has shown no significant visible or stress related effects with no significant increase in N status.     

Can the foliar nitrogen concentration of upland vegetation be used for predicting atmospheric nitrogen deposition? Evidence from field surveys

The deposition of atmospheric nitrogen can be enhanced at high altitude sites as a consequence of cloud droplet deposition and orographic enhancement of wet deposition on hills. The degree to which the increased deposition of nitrogen influences foliar nitrogen concentration in a range of upland plant species was studied in a series of field surveys in northern Britain. A range of upland plant species sampled along altitudinal transects at sites of known atmospheric nitrogen deposition showed marked increases in foliar nitrogen concentration with increasing nitrogen deposition and altitude (and hence with decreasing temperature). For Nardus stricta L., Deschampsia flexuosa (L.) Trin., Calluna vulgaris (L.) Hull, Erica cinerea L. and Hylocomium splendens (Hedw.) Br. Eur. on an unpolluted hill, foliar nitrogen increased by 0.07, 0.12, 0.15, 0.08 and 0.04% dry weight respectively for each 1 kg ha(-1) year(-1) increase in nitrogen deposition. Most species showed an approximately linear relationship between foliar nitrogen concentration and altitude but no trend with altitude for foliar phosphorus concentration. This provided evidence that the tissue nutrient status of upland plants reflects nutrient availability rather than the direct effects of climate on growth. However, differences in the relationship between foliar nitrogen concentration and atmospheric nitrogen deposition for N. stricta sampled on hills in contrasting pollution climates show that the possibility of temperature-mediated growth effects on foliar nitrogen concentration should not be ignored. Thus, there is potential to use upland plant species as biomonitors of nitrogen deposition, but the response of different species to nitrogen addition, in combination with climatic effects on growth, must be well characterised.     

Fluxes of oxidised and reduced nitrogen above a mixed coniferous forest exposed to various nitrogen emission sources

Concentrations of nitrogen gases (NH(3), NO(2), NO, HONO and HNO(3)) and particles (pNH(4) and pNO(3)) were measured over a mixed coniferous forest impacted by high nitrogen loads. Nitrogen dioxide (NO(2)) represented the main nitrogen form, followed by nitric oxide (NO) and ammonia (NH(3)). A combination of gradient method (NH(3) and NO(x)) and resistance modelling techniques (HNO(3), HONO, pNH(4) and pNO(3)) was used to calculate dry deposition of nitrogen compounds. Net flux of NH(3) amounted to -64 ng N m(-2) s(-1) over the measuring period. Net fluxes of NO(x) were upward (8.5 ng N m(-2) s(-1)) with highest emission in the morning. Fluxes of other gases or aerosols substantially contributed to dry deposition. Total nitrogen deposition was estimated at -48 kg N ha(-1) yr(-1) and consisted for almost 80% of NH(x). Comparison of throughfall nitrogen with total deposition suggested substantial uptake of reduced N (+/-15 kg N ha(-1) yr(-1)) within the canopy.     

Effects of nitrogen with and without acidified sulphur on an ectomycorrhizal community in a Sitka spruce (Picea sitchensis Bong. Carr) forest

This preliminary study investigated the effects of enhanced nitrogen (NH4NO3 at 48 kg ha(-1) y(-1)), sulphur (Na2SO4 at 50 kg ha(-1) y(-1)), acidified nitrogen and sulphur (H2SO4 + NH4NO3) at pre-stated doses (pH 2.5), and acidified nitrogen and sulphur deposition at double these doses on the ectomycorrhizal community associated with a 13-year-old Sitka spruce (Picea sitchensis) forest. Sulphur deposition had little impact on below ground ectomycorrhizal diversity, but stimulated sporocarp production. Nitrogen inputs increased below ground colonisation compared to acidified nitrogen and sulphur, largely due to an increase in Tylospora fibrillosa colonisation. Sporocarp production and ectomycorrhizal root colonisation by Lactarius rufus were reduced in the nitrogen treated plots. These observations suggest that nitrogen deposition to a young plantation may suppress ectomycorrhizal fungi producing large sporocarps. It is proposed that enhanced nitrogen deposition increases ectomycorrhizal nitrogen assimilation, consuming more carbon and leaving less for extrametrical mycelium and sporocarp development.     

Critical loads and their exceedances at intensive forest monitoring sites in Europe

Intensive forest monitoring by means of harmonised methods has been conducted in Europe for more than a decade. Risks of atmospheric nitrogen and sulphur deposition are assessed by means of calculations of critical loads and their exceedances. In the present study throughfall and bulk deposition of nitrate (N-NO(3)), ammonium (N-NH(4)) and sulphate (S-SO(4)) show marked spatial patterns and temporal trends. In the period of observation (1999-2004), sulphate deposition on intensive monitoring plots decreased by about one quarter. This is in line with the reduction of S deposition by 70% since 1981 in Europe as a result of successful air pollution control politics under the Convention on Long-range Transboundary Air Pollution (CLRTAP). However, sulphate and especially nitrate and ammonium deposition were found to still exceed critical loads at many forest sites, indicating a continued need for further implementation of air pollution abatement strategies.     

Deposition of atmospheric ammonia to moorlands

Micrometeorological methods were applied to measure fluxes of atmospheric ammonia (NH3) to moorlands. Measurements were made in a wide variety of surface conditions and included both Calluna vulgaris (L.) Hull and Eriophorum vaginatum L. dominated sites. NH3 was found to deposit rapidly to all the sites investigated, providing large deposition velocities (Vd, typically 10-40 mm s(-1)) and usually minimal surface resistances (rc). A small number of measurements were made in frozen conditions and suggest a possible exception to this pattern with mean rc of 50-200 s m(-1). The effect of vegetation drying was also investigated and a possible increase in rc observed, though this was small (< 10 s m(-1)). The results are interpreted in terms of the processes controlling exchange; it is shown that NH3 deposition is predominantly to the leaf surfaces and that the net NH3 compensation point approaches zero. Annual estimates show that dry deposition of NH3 is a major source of atmospheric nitrogen to moorland ecosystems. For two typical UK sites subject to background air concentrations, NH3 dry deposition is of similar magnitude to equivalent NH4+ inputs in wet deposition. In the vicinity of emission sources, NH3 dry deposition is expected to dominate inputs of atmospheric nitrogen.     

Crown condition of Norway spruce in relation to sulphur and nitrogen deposition and soil properties in southeast Norway

Atmospheric deposition of sulphur and nitrogen compounds may lead to enhanced leaching of base cations, accumulation of nitrogen in organic matter, lowered pH and increased concentration of toxic aluminium in soil, which in turn may affect the vitality of forest trees. A general monitoring of forest condition has been initiated in many European countries, partly in order to reveal stresses caused by acidification. However, forest condition is also affected by many other factors. This paper examines a seven-year series of crown-condition data from Local County Monitoring Plots in Norway spruce stands in Norway. Average, time trend and lability variables were calculated for crown density and crown colour for each plot. Wet deposition of sulphate, ammonium and nitrate for each plot were estimated using data from the national air and precipitation monitoring programmes. Soil data are based on soil sampling within the plots. The analysis gave no evident support for the hypothesized negative effect on crown condition from sulphur and nitrogen deposition and related alterations in soil.     

Changes in the atmospheric deposition of acidifying compounds in the UK between 1986 and 2001

Emissions of a precursor of acidity in precipitation, sulphur dioxide (SO2), declined in the UK and the EU (15) by 71% and 72%, respectively, between 1986 and 2001, while nitrous oxide emissions declined by about 40%. Acidity in UK precipitation and the deposition of sulphate in precipitation halved during this period, but reductions were larger in the English Midlands than at the west coast and in high rainfall areas (>2000 mm). There is evidence that the smaller reductions in sulphur deposition in the west and south are due in part to shipping sources of SO2. Reductions in sulphur dry deposition (74%) are larger than in wet deposition (45%), due to changes in the canopy resistance to dry deposition. For reduced nitrogen, there has been a small (10%) reduction in emissions and deposition, while for oxidized nitrogen, a substantial reduction in emissions (40%) occurred but wet deposition of nitrate changed by less than 10%.     

Assessing airborne pollution effects on bryophytes: lessons learned through long-term integrated monitoring in Austria

The study uses measured and calculated data on airborne pollutants, particularly nitrogen (ranges between 28 to 43kgN*ha(-1)*yr(-1)) and sulphur (10 to 18kgSO(4)-S*ha(-1)*yr(-1)), in order to assess their long-term (1992 to 2005) effects on bryophytes at the UN-ECE Integrated Monitoring site 'Z?belboden' in Austria. Bryophytes were used as reaction indicators on 20 epiphytic plots using the IM monitoring method and on 14 terrestrial plots using standardised photography. The plots were recorded in the years 1992, 1993, 1998, and 2004/2005. Most species remained stable in terms of their overall population size during the observed period, even though there were rapid turnover rates of a large percentage of species on all investigated plots. Only a few bryophytes (Hypnum cupressiforme, Leucodon sciuroides) responded unambiguously to N and S deposition. Nitrogen deposition had a weak but significant effect on the distribution of bryophyte communities. However, the time shifts in bryophyte communities did not depend on total deposition of N and S.