Simon J.M. Caporn

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.

Evidence that soil carbon pool determines susceptibility of semi-natural ecosystems to elevated nitrogen leaching

Deposition of reactive nitrogen (N) compounds has the potential to cause severe damage to sensitive soils and waters, but the process of ‘nitrogen saturation’ is difficult to demonstrate or predict. This study compares outputs from a simple carbon–nitrogen model with observations of (1) regional- and catchment-scale relationships between surface water nitrate and dissolved organic carbon (DOC), as an indicator of catchment carbon (C) pool; (2) inter-regional variations in soil C/N ratios; and (3) plot scale soil and leachate response to long-term N additions, for a range of UK moorlands. Results suggest that the simple model applied can effectively reproduce observed patterns, and that organic soil C stores provide a critical control on catchment susceptibility to enhanced N leaching, leading to high spatial variability in the extent and severity of current damage within regions of relatively uniform deposition. Results also support the hypothesis that the N richness of organic soils, expressed as C/N ratio, provides an effective indicator of soil susceptibility to enhanced N leaching. The extent to which current C/N is influenced by N deposition, as opposed to factors such as climate and vegetation type, cannot be unequivocally determined on the basis of spatial data. However, N addition experiments at moorland sites have shown a reduction in organic soil C/N. A full understanding of the mechanisms of N-enrichment of soils and waters is essential to the assessment of current sensitivity to, and prediction of future damage from, globally increasing reactive nitrogen deposition.

The Role of Nitrogen Deposition in Widespread Plant Community Change Across Semi-natural Habitats

Experimental studies have shown that deposition of reactive nitrogen is an important driver of plant community change, however, most of these experiments are of short duration with unrealistic treatments, and conducted in regions with elevated ambient deposition. Studies of spatial gradients of pollution can complement experimental data and indicate whether the potential impacts demonstrated by experiments are actually occurring in the ‘real world’. However, targeted surveys exist for only a very few habitats and are not readily comparable. In a coordinated campaign, we determined the species richness and plant community composition of five widespread, semi-natural habitats across Great Britain in sites stratified along gradients of climate and pollution, and related these ecological parameters to major drivers of biodiversity, including climate, pollution deposition, and local edaphic factors. In every habitat, we found reduced species richness and changed species composition associated with higher nitrogen deposition, with remarkable consistency in relative species loss across ecosystem types. Whereas the diversity of mosses, lichens, forbs, and graminoids declines with N deposition in different habitats, the cover of graminoids generally increases. Considered alongside previous experimental studies and survey work, our results provide a compelling argument that nitrogen deposition is a widespread and pervasive threat to terrestrial ecosystems.

The effect of long-term nitrogen additions on the bryophyte cover of upland acidic grasslands

Abstract Regular applications of NH4NO3 (3.5–14 g N m−2 yr−l) and (NH4)2SO4 (14 g N m−2 yr−l) to areas of acidic grassland in the Derbyshire Peak District over a period of six years have resulted in marked changes in the abundance of the bryophyte species present on the site. A dose-related reduction in bryophyte cover, significant at all levels of nitrogen addition, was obtained after only one year of applications and this effect has been maintained over a further five years of treatment. (NH4)2SO4 additions at the same rate as the highest NH4NO3 input (14 g N m−2 yr−1) produced greater reductions in cover with losses of 80–95% compared with 45–55% for the 14 g NH4NO3 treatment. Soil pH measurements taken from cores collected in December 1994 showed a small reduction in the pH of the (NH4)2SO4 treated plots (4.03 compared with 4.41 for the controls), whereas the NH4NO3 treatments were unaffected. The mean stem density of the dominant bryophyte species Rhytidiadelphus squarrosus was significantly reduced at the lowest level of nitrogen addition and this was associated with marked increases in the total stem nitrogen content of this species. Pleurozium schreberi, present at much lower cover values, however showed no significant change except at the highest level of application, suggesting differential effects of the treatments on these two species. These results are indicative of significant losses in the bryophyte cover of acidic grasslands at atmospheric input rates of 5 g N m−2 yr−1 or less, and suggest that these species may be particularly at risk from the high atmospheric nitrogen deposition rates that are becoming a feature of many upland areas.

Nitrogen Deposition Reduces Plant Diversity and Alters Ecosystem Functioning: Field-Scale Evidence from a Nationwide Survey of UK Heathlands

Findings from nitrogen (N) manipulation studies have provided strong evidence of the detrimental impacts of elevated N deposition on the structure and functioning of heathland ecosystems. Few studies, however, have sought to establish whether experimentally observed responses are also apparent under natural, field conditions. This paper presents the findings of a nationwide field-scale evaluation of British heathlands, across broad geographical, climatic and pollution gradients. Fifty two heathlands were selected across an N deposition gradient of 5.9 to 32.4 kg ha−1 yr−1. The diversity and abundance of higher and lower plants and a suite of biogeochemical measures were evaluated in relation to climate and N deposition indices. Plant species richness declined with increasing temperature and N deposition, and the abundance of nitrophilous species increased with increasing N. Relationships were broadly similar between upland and lowland sites, with the biggest reductions in species number associated with increasing N inputs at the low end of the deposition range. Both oxidised and reduced forms of N were associated with species declines, although reduced N appears to be a stronger driver of species loss at the functional group level. Plant and soil biochemical indices were related to temperature, rainfall and N deposition. Litter C:N ratios and enzyme (phenol-oxidase and phosphomonoesterase) activities had the strongest relationships with site N inputs and appear to represent reliable field indicators of N deposition. This study provides strong, field-scale evidence of links between N deposition - in both oxidised and reduced forms - and widespread changes in the composition, diversity and functioning of British heathlands. The similarity of relationships between upland and lowland environments, across broad spatial and climatic gradients, highlights the ubiquity of relationships with N, and suggests that N deposition is contributing to biodiversity loss and changes in ecosystem functioning across European heathlands.

The effect of exposure to NO2 and SO2 on frost hardiness in Calluna vulgaris

Heather (Calluna vulgaris) was grown in pots in a natural heathland soil and exposed in outdoor fumigation chambers (‘solardomes’) to 40 nl l−1 of both nitrogen dioxide (NO2) and sulphur dioxide (SO2) simultaneously. Plants exposed to NO2 and SO2 for 8 months over a growing season (February–November) showed increases in the growth of shoots (+37%) and the whole plant (+15%) and a raised ratio of root to shoot dry matter in comparison with control (charcoal-filtered air) plants. Fumigation raised the average foliar concentrations of nitrogen (+34%) and sulphur (+173%). The improvements in growth due to pollutant exposure were countered by reductions in tolerance to experimental acute frost. Cellular damage of heather shoots was assessed using measurements of electrolyte leakage from cut shoots following controlled over-night frosts. The rates of leakage were consistently increased in those plants that had been exposed to NO2 and SO2 for 5 months or more in comparison with non-fumigated controls. In some cases a greater leakage rate was recorded in fumigated plants than in controls even in the absence of freezing temperatures. The pollutants caused a similar reduction in frost tolerance whether exposure was given during the hardening period (August–January) or the de-hardening stage (November–April). These results support the hypothesis that low concentrations of air pollutants can reduce the tolerance of plants to freezing stress.

Bio-indicators of nitrogen pollution in heather moorland

Heather moorlands are internationally important ecosystems that are highly sensitive to eutrophication and acidification by reactive atmospheric nitrogen (N) deposition. We used a long-term experiment simulating wet-deposition of N on heather moorland to identify potential bio-indicators of N deposition. These indicators were subsequently employed in a survey covering a N deposition gradient ranging from approximately 7 to 31kg N ha(-1) yr(-1), at selected sites throughout the UK. In this regional survey litter phenol oxidase activity and bryophyte species richness were negatively associated with N deposition. Calluna vulgaris N:P ratios and litter extractable N were positively correlated with N deposition. The use of the suite of four bio-indicators has the potential to provide rapid assessment of the extent of N saturation of heather moorland sites and moorland ecosystem functioning, and has significant advantages over reliance on single measures such as soil N status or an individual bio-indicator species.

Bacterial and Fungal Communities in a Degraded Ombrotrophic Peatland Undergoing Natural and Managed Re-Vegetation

The UK hosts 15–19% of global upland ombrotrophic (rain fed) peatlands that are estimated to store 3.2 billion tonnes of carbon and represent a critical upland habitat with regard to biodiversity and ecosystem services provision. Net production is dependent on an imbalance between growth of peat-forming Sphagnum mosses and microbial decomposition by microorganisms that are limited by cold, acidic, and anaerobic conditions. In the Southern Pennines, land-use change, drainage, and over 200 years of anthropogenic N and heavy metal deposition have contributed to severe peatland degradation manifested as a loss of vegetation leaving bare peat susceptible to erosion and deep gullying. A restoration programme designed to regain peat hydrology, stability and functionality has involved re-vegetation through nurse grass, dwarf shrub and Sphagnum re-introduction. Our aim was to characterise bacterial and fungal communities, via high-throughput rRNA gene sequencing, in the surface acrotelm/mesotelm of degraded bare peat, long-term stable vegetated peat, and natural and managed restorations. Compared to long-term vegetated areas the bare peat microbiome had significantly higher levels of oligotrophic marker phyla (Acidobacteria, Verrucomicrobia, TM6) and lower Bacteroidetes and Actinobacteria, together with much higher ligninolytic Basidiomycota. Fewer distinct microbial sequences and significantly fewer cultivable microbes were detected in bare peat compared to other areas. Microbial community structure was linked to restoration activity and correlated with soil edaphic variables (e.g. moisture and heavy metals). Although rapid community changes were evident following restoration activity, restored bare peat did not approach a similar microbial community structure to non-eroded areas even after 25 years, which may be related to the stabilisation of historic deposited heavy metals pollution in long-term stable areas. These primary findings are discussed in relation to bare peat oligotrophy, re-vegetation recalcitrance, rhizosphere-microbe-soil interactions, C, N and P cycling, trajectory of restoration, and ecosystem service implications for peatland restoration.

The legacy of nitrogen pollution in heather moorlands: Ecosystem response to simulated decline in nitrogen deposition over seven years

Eutrophication and acidification of heather moorlands by chronic atmospheric nitrogen (N) pollution, is of major concern within these internationally important ecosystems. However, in the UK and Western Europe generally emissions of NO(y) and NH(x) peaked during the 20th century. Due to the history and scale of atmospheric N pollution, the legacy of these high levels of N deposition, through accumulation in soil, may hinder or prevent ecosystem recovery. Effects of N pollution on heather moorland were investigated throughout the ecosystem including; the dominant plant species, Calluna vulgaris, the bryophyte and lichen community and the soil system using a long-term experiment simulating wet N deposition. We observed an increase in C. vulgaris height, shoot extension and canopy density, litter mineral N, total N concentration, N:P and C:N ratios in response to N addition. Bryophyte species diversity, bryophyte and lichen frequency and the frequency of two individual bryophyte species (Lophozia ventricosa and Campylopus flexuosus) were significantly reduced by N addition. We developed an N recovery experiment, using a split-plot design, on the long-term N treatment plots to investigate ecosystem response to a simulated decline in N deposition. Two years after cessation of N treatment the only ecosystem component that responded to the recovery experiment was C. vulgaris shoot extension, however after seven years of recovery there were significant declines in litter total N concentration and mineral N and an increase in litter C:N ratio. Although bryophytes and lichens form a close relationship with atmospheric N deposition these organisms did not show a significant response to the N recovery experiment, two years after cessation of N treatment. These data indicate that low nutrient ecosystems, such as moorlands, have the capacity to respond to declines in N deposition however the accumulation of pollution may hinder recovery of sensitive organisms, such as bryophytes and lichens.

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 its 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.