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.     

Effects of simultaneous ozone exposure and nitrogen loads on carbohydrate concentrations, biomass, growth, and nutrient concentrations of young beech trees (Fagus sylvatica)

Beech seedlings were grown under different nitrogen fertilisation regimes (0, 20, 40, and 80 kg Nha(-1)yr(-1)) for three years and were fumigated with either charcoal-filtered (F) or ambient air (O3). Nitrogen fertilisation increased leaf necroses, aphid infestations, and nutrient ratios in the leaves (N:P and N:K), as a result of decreased phosphorus and potassium concentrations. For plant growth, biomass accumulation, and starch concentrations, a positive nitrogen effect was found, but only for fertilisations of up to 40 kg Nha(-1) yr(-1). The highest nitrogen load, however, reduced leaf area, leaf water content, growth, biomass accumulation, and starch concentrations, whereas soluble carbohydrate concentrations were enhanced. The ozone fumigation resulted in reduced leaf area, leaf water content, shoot growth, root biomass accumulation, and decreased starch, phosphorus, and potassium concentrations, increasing the N:P and N:K ratios. A combined effect of the two pollutants was detected for the leaf area and the shoot elongation, where ozone fumigation amplified the nitrogen effects.     

Nitrogen dynamics of a mountain forest on dolomitic limestone--a scenario-based risk assessment

The dominant nitrogen (N) fluxes were simulated in a mountain forest ecosystem on dolomitic bedrock in the Austrian Alps. Based on an existing small-scale climate model the simulation encompassed the present situation and a 50-yr projection. The investigated scenarios were current climate, current N deposition (SC1) and future climate (+2.5 degrees C and +10% annual precipitation) with three levels of N deposition (SC2, 3, 4). The microbially mediated N transformation, including the emission of nitrogen oxides, was calculated with PnET-N-DNDC. Soil hydrology was calculated with HYDRUS and was used to estimate the leaching of nitrate. The expected change of the forest ecosystem due to changes of the climate and the N availability was simulated with PICUS. The incentive for the project was the fact that forests on dolomitic limestone stock on shallow Rendzic Leptosols that are rich in soil organic matter are considered highly sensitive to the expected environmental changes. The simulation results showed a strong effect due to increased temperatures and to elevated levels of N deposition. The outflux of N, both as nitrate (6-25kg Nha(-1)yr(-1)) and nitrogen oxides (1-2kg Nha(-1)yr(-1)), from the forest ecosystem are expected to increase. Temperature exerts a stronger effect on the N(2)O emission than the increased rate of N deposition. The main part of the N emission will occur as N(2) (15kg Nha(-1)yr(-1)). The total N loss is partially offset by increased rates of N uptake in the biomass due to an increase in forest productivity.     

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.     

Effects of increased deposition of atmospheric nitrogen on an upland Calluna moor: N and P transformations

This study determined the effects of increased N deposition on rates of N and P transformations in an upland moor. The litter layer and the surface of the organic Oh horizon were taken from plots that had received long-term additions of ammonium nitrate at rates of 40, 80 and 120 kg N ha(-1) yr(-1). Net mineralisation processes were measured in both field and laboratory incubations. Soil phosphomonoesterase (PME) activity and rates of N(2)O release were measured in laboratory incubations and root-surface PME activity measured in laboratory microcosms using Calluna vulgaris bioassay seedlings. Net mineralisation rates were relatively slow, with net ammonification consistently stimulated by N addition. Net nitrification was marginally stimulated by N addition in the laboratory incubation. N additions also increased soil and root-surface (PME) activity and rates of N(2)O release. Linear correlations were found between litter C:N ratio and all the above processes except net nitrification in field incubations. When compared with data from a survey of European forest sites, values of litter C:N ratio were greater than a threshold below which substantial, N input-related increases in net nitrification rates occurred. The maintenance of high C:N ratios with negligible rates of net nitrification was associated with the common presence of ericaceous litter and a mor humus layer in both this moorland as well as the forest sites.     

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.     

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.     

Interactive effects of nitrogen deposition and fire on plant and soil chemistry in an alpine heathland

The response of alpine heathland vegetation and soil chemistry to N additions of 0, 10, 20 and 50 kg N ha⁻¹ year⁻¹ in combination with simulated accidental fire (+/−) was monitored over a 5-year period. N addition caused rapid and significant increases in plant tissue N content and N:P and N:K of Calluna vulgaris, suggesting increasing phosphorus and potassium limitation of growth. Soil C:N declined significantly with N addition, indicating N saturation and increasing likelihood of N leakage. Fire further decreased soil C:N and reduced potential for sequestration of additional N. This study shows that alpine heathlands, which occupy the headwaters of many rivers, have limited potential to retain deposited N and may rapidly become N saturated, leaking N into downstream communities and surface waters.     

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.     

Phosphorus quantity-intensity relationships and agronomic measures of P in surface layers of soil from a long-term slurry experiment

Soils from a long-term slurry experiment established in 1970 at Hillsborough, Northern Ireland, were used in the experiment. The site has a clay loam soil overlying Silurian shale. Seven treatments were used with three replicate plots per treatment. Control plots were treated with mineral fertiliser supplying 200 kg N, 32 kg P and 160 kg K ha(-1) yr(-1). Slurry treatment plots were in two blocks and treated with either pig or cow slurry supplied at 50, 100 or 200 m3 ha(-1) yr(-1). Agronomic measures of P determined on 10-cm soil cores were compared with measured P quantity/intensity (Q/I) parameters from fitted sorption and desorption isotherms. Phosphorus affinity constant was found to be significantly and negatively correlated with P loading of soils. Desorption rate coefficient also increased significantly with increase in P loading from slurry, although there was no significant difference between slurry types (cow vs. pig). In contrast, while agronomic measures of P (water-soluble P, Olsen P, calcium chloride-extractable P, degree of P saturation (DPS)) also correlated significantly with P loading and total P (TP) in the soils, there was a separation and significant differences between the cow and pig slurry treatment blocks, with the former being much lower. Phosphorus inputs to pig slurry treated plots were much higher than to equivalent cow slurry plots over the first 15 years of the study but declined sharply over the most recent 10 years to more or less par. Conventional measures of agronomic P such as Olsen P and DPS, measure only P accumulation over the longer term and indicated only the higher content of P accumulating in soil of pig slurry treatments. Risk of P loss estimated by Q/I parameters appeared to show very similar behaviour between the two slurry types in line with more recent manurial additions but in contradiction of P accumulation statistics.     

The influence of nitrogen deposition,competition and desiccation on growth and regeneration of Racomitrium lanuginosum (Hedw.) brid

Racomitrium lanuginosum shoot growth was studied under the combined effects of N deposition (0, 20, 40 and 60 kg N ha(-1) year(-1)). competition with Festuca ovina, and a drought pre-treatment. Moss regeneration from shoot fragments was also investigated. Growth was initially stimulated at the 60 kg N level. However, after 6 months, growth was lower in all N treatments than in the 0 kg N control. Reductions in shoot growth first became apparent in the pre-desiccated moss, while moss shoots grew longer when surrounded by a F. ovina canopy. Optimum regeneration occurred at 20-40 kg N on bare soil, and at 0-20 kg N under a F. ovina canopy. These results suggest that current N deposition in upland Wales is already detrimental to growth of this species, and to regeneration under certain conditions. This species may be affected under predicted climatic scenarios of increased summer drought in Britain.     

The interactions between plant growth,vegetation structure and soil processes in semi-natural acidic and calcareous grasslands receiving long-term inputs of simulated pollutant nitrogen deposition

Regular applications of ammonium nitrate (35-140 kg N ha(-1) year(-1)) and ammonium sulphate (140 kg N ha(-1) year(-1)) to areas of acidic and calcareous grassland in the Derbyshire Peak District over a period of 6 years, have resulted in significant losses in both overall plant cover, and the abundance of individual species, associated with clear and dose-related increases in shoot nitrogen content. No overall growth response to nitrogen treatment was seen at any stage in the experiment. Phosphorus additions to the calcareous plots did however lead to significant increases in plant cover and total biomass, indicative of phosphorus limitation in this system. Clear and dose-related increases in soil nitrogen mineralization rates were also obtained, consistent with marked effects of the nitrogen additions on soil processes. High nitrification rates were seen on the calcareous plots, and this process was associated with significant acidification of the 140 kg N ha(-1) year(-1) treatments.     

Effects of increased deposition of atmospheric nitrogen on an upland moor: leaching of N species and soil solution chemistry

This study was designed to investigate the leaching response of an upland moorland to long-term (10 yr) ammonium nitrate additions of 40, 80 and 120 kg N ha(-1) yr(-1) and to relate this response to other indications of potential system damage, such as acidification and cation displacement. Results showed increases in nitrate leaching only in response to high rates of N input, in excess of 96 and 136 kg total N input ha(-1) yr(-1) for the organic Oh horizon and mineral Eag horizon, respectively. Individual N additions did not alter ammonium leaching from either horizon and ammonium was completely retained by the mineral horizon. Leaching of dissolved organic nitrogen (DON) from the Oh horizon was increased by the addition of 40 kg N ha(-1) yr(-1), but in spite of increases, retention of total dissolved nitrogen reached a maximum of 92% and 95% of 80 kg added N ha(-1) yr(-1) in the Oh and Eag horizons, respectively. Calcium concentrations and calcium/aluminium ratios were decreased in the Eag horizon solution with significant acidification mainly in the Oh horizon leachate. Nitrate leaching is currently regarded as an early indication of N saturation in forest systems. Litter C:N ratios were significantly lowered but values remained above a threshold predicted to increase leaching of N in forests.     

Agronomic measures of P, Q/I parameters and lysimeter-collectable P in subsurface soil horizons of a long-term slurry experiment

Soils from a long-term slurry experiment established in 1970 at Hillsborough, Northern Ireland, were used in the experiment. The site has a clay loam soil overlying Silurian shale. Seven treatments were used with three replicate plots per treatment under the following manurial regimes: (1) mineral fertiliser supplying 200 kg N, 32 kg P and 160 kg K ha(-1) yr(-1); (2)-(4) pig slurry applied at 50, 100 or 200 m3 ha(-1) yr(-1); (5)-(7) cow slurry applied at 50, 100 or 200 m3 ha(-1) yr(-1). Agronomic measures of P determined on subsurface layers down to 90 cm were compared with sorption isotherm data and rates of desorption. Adsorption isotherms were fitted using a standard Langmuir model. Data were compared with soluble (molybdate-reactive) P levels in soil water collected at 35 and 90 cm using PTFE suction cup lysimeters. Agronomically available P was concentrated in the top 30 cm of soil in all treatments. The accumulation of P in surface layers of the plots was significantly greater in the pig slurry treatments compared to the cow slurry, reflecting the history of P amendments. Nevertheless, over a period of a year, molybdate-reactive phosphorus (MRP) concentrations in lysimeter collections was consistently higher at 35 cm depth in the highest cow slurry treatment (7) compared to the equivalent pig slurry treatment (4). Either the movement of soluble P down the profile is facilitated by the higher organic content of cow slurry or P movement is not directly related to P accumulation in the soils. In addition, it is hypothesised that P movement down the soil profile depends upon two separate mechanisms. First, a 'break' point above which the accumulated P in the surface horizons is less strongly held and therefore amenable to dissolution and movement down the profile. Second, a mechanism by which some solute P from the surface horizons can travel rapidly through horizons of low P status to greater depth in the soil, i.e., by preferential flow.     

Effect of amendment C:N ratio on plant richness, cover and metal content for acidic Pb and Zn mine tailings in Leadville, Colorado

Biosolids and woody debris were applied with target C:N ratios of 8:1 to 50:1 to phytotoxic, acidic, high metal mine tailings to test the effect of amendment C:N ratio on native plant restoration. Total soil C decreased over time indicating an active microbial community. The 8:1 treatment initially had no growth, the highest plant cover for the final sampling (86.8+/-13.8%) and the lowest number of species (3.33+/-0.4). The greatest number of species was in the 30:1 treatment (5.44+/-0.45). Plant cover increased over time for all treatments from 44.7% in 2001 to 71% in 2005. This response was consistent across all except for the 30:1 treatment, which showed a slight decrease in the final year (65+/-11%). Volunteer species and evidence of animal grazing were observed in all amended plots. Results indicate that a C:N ratio>/=20:1 increased species diversity.     

Base cation depletion, eutrophication and acidification of species-rich grasslands in response to long-term simulated nitrogen deposition

Pollutant nitrogen deposition effects on soil and foliar element concentrations were investigated in acidic and limestone grasslands, located in one of the most nitrogen and acid rain polluted regions of the UK, using plots treated for 8-10years with 35-140kg Nha(-2)y(-1) as NH(4)NO(3). Historic data suggests both grasslands have acidified over the past 50years. Nitrogen deposition treatments caused the grassland soils to lose 23-35% of their total available bases (Ca, Mg, K, and Na) and they became acidified by 0.2-0.4 pH units. Aluminium, iron and manganese were mobilised and taken up by limestone grassland forbs and were translocated down the acid grassland soil. Mineral nitrogen availability increased in both grasslands and many species showed foliar N enrichment. This study provides the first definitive evidence that nitrogen deposition depletes base cations from grassland soils. The resulting acidification, metal mobilisation and eutrophication are implicated in driving floristic changes.     

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.     

Winter runoff losses of phosphorus from paddy soils in the Taihu Lake Region of South China

A winter wheat field plot experiment was conducted on two types of paddy soils, from November, 2000 to June, 2001 to assess P losses to its surrounding watercourses by runoff in the Taihu Lake Region. Commercial NPK compound fertilizer and single superphosphate fertilizer were applied to furnish 0, 20, 80, and 160 kg P ha(-1). The experiments consisted of six replicates of each treatment in Changshu site and four replicates in Anzhen site, with a plot size of 5x6 m2 in a randomized block design. Results revealed that the average concentration of dissolved P (DP), particulate P (PP), and total P (TP) in runoff water during the winter season was 0.13, 0.90 and 1.04 mg P l(-1) respectively, from P20 plots in Anzhen site. While it was 0.67, 1.08 and 1.75 mg P l(-1) respectively, from P20 plots in Changshu site. The seasonal TP load (mass loss) from P20 plot ranged from a low of 290.88 g P ha(-1)season(-1) to a high of 483.54 g P ha(-1)season(-1), with a mean of 382.29 g P ha(-1)season(-1) in Anzhen, but from 444.92 to 752.21 g P ha(-1)season(-1), with a mean of 539.13 g P ha(-1)season(-1) in Changshu. Both in Anzhen and Changshu PP represented a major portion of the TP lost in runoff, the average PP/TP was about more than 80% in P0 and P20 plot, but it was decreased with the increase of P rate. The average seasonal P loads (DP, PP, and TP) in Changshu were greater than in Anzhen although runoff volume in Anzhen (45 mm season(-1)) was more than in Changshu (36 mm season(-1)). This was probably associated with the differences of soil physical and chemical properties between the two sites. Phosphate fertilizer rate significantly affected P concentrations and P loads by runoff. Both the mean concentrations and the average seasonal P loads from the P80 plots were lower than from the P160 plots, but obviously higher than from the P20 and P0 plots. There was no significant difference found between the P20 plots and the P0 plots both in Anzhen and Changshu sites. It indicated that P loads by runoff would be greatly increased in 5-10 years due to the accumulation of soil P if 20 kg P ha(-1) applied each wheat season in this area.     

Effects of nitrogen deposition and soil fertility on cover and physiology of Cladonia foliacea (Huds.) Willd., a lichen of biological soil crusts from Mediterranean Spain

We are fertilizing a thicket with 0, 10, 20 and 50 kg nitrogen (N) ha⁻¹ yr⁻¹ in central Spain. Here we report changes in cover, pigments, pigment ratios and FvFm of the N-tolerant, terricolous, lichen Cladonia foliacea after 1-2 y adding N in order to study its potential as biomarker of atmospheric pollution. Cover tended to increase. Pigments increased with fertilization independently of the dose supplied but only significantly with soil nitrate as covariate. β-carotene/chlorophylls increased with 20-50 kg N ha⁻¹ yr⁻¹ (over the background) and neoxanthin/chlorophylls also increased with N. (Neoxanthin+lutein)/carotene decreased with N when nitrate and pH seasonalities were used as covariates. FvFm showed a critical load above 40 kg N ha⁻¹ yr⁻¹. Water-stress, iron and copper also explained variables of lichen physiology. We conclude that this tolerant lichen could be used as biomarker and that responses to N are complex in heterogeneous Mediterranean-type landscapes.