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.

What is the most ecologically-meaningful metric of nitrogen deposition?

Nitrogen (N) deposition poses a severe risk to global terrestrial ecosystems, and managing this threat is an important focus for air pollution science and policy. To understand and manage the impacts of N deposition, we need metrics which accurately reflect N deposition pressure on the environment, and are responsive to changes in both N deposition and its impacts over time. In the UK, the metric typically used is a measure of total N deposition over 1-3 years, despite evidence that N accumulates in many ecosystems and impacts from low-level exposure can take considerable time to develop. Improvements in N deposition modelling now allow the development of metrics which incorporate the long-term history of pollution, as well as current exposure. Here we test the potential of alternative N deposition metrics to explain vegetation compositional variability in British semi-natural habitats. We assembled 36 individual datasets representing 48,332 occurrence records in 5479 quadrats from 1683 sites, and used redundancy analyses to test the explanatory power of 33 alternative N metrics based on national pollutant deposition models. We find convincing evidence for N deposition impacts across datasets and habitats, even when accounting for other large-scale drivers of vegetation change. Metrics that incorporate long-term N deposition trajectories consistently explain greater compositional variance than 1-3 year N deposition. There is considerable variability in results across habitats and between similar metrics, but overall we propose that a thirty-year moving window of cumulative deposition is optimal to represent impacts on plant communities for application in science, policy and management.

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.

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.

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.

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.

Crystal occurrence and wax disruption on leaf surfaces of cabbage treated with simulated acid rain.

Epicuticular waxes on leaves of Brassica oleracea L. (cabbage) were studied using scanning electron microscopy after a single treatment with simulated rain of pH 5.6, 3.0 or 2.5 which was either sprayed on to plants in an exposure chamber or applied as droplets with a micropipette. Treatments with acidified rain caused serious structural degradation of the wax crystals. The alteration of crystalline wax structures was similar for leaves treated with nitric acid solutions, but less severe, than for leaves treated with sulphuric acid solutions. With both H2 SO4 , and HNOa -derived rain solutions numerous gypsum (CaSO4 ) crystals were found in and near lesions on the leaves treated with rain of pH 3.0 and 2.5. The crystals probably resulted from damage of cuticular membranes by acidic rain which significantly altered their permeability to ions in the area of lesions. Crystalline leaf waxes may be an important target for acidic pollutants, and the physiological consequences of their degradation are discussed.

The effects of oxides of nitrogen and carbon dioxide enrichment on photosynthesis and growth of lettuce (Lactuca sativa L.).

The response of glasshouse crops to the nitrogen oxide pollutants which may be generated during enrichment with O2 has been studied in controlled environments. Lettuce (Lactuca sativa L. cv. Ambassador) was grown in air containing either low CO2 , (380µmol mol-1 ), high CO2 , (1200 µmol mol -1 ), or high CO2 plus oxides of nitrogen? (NOx )- Carbondioxide enrichment increased the rate of emergence and expansion of leaves and the growth of Young plants. Addition of NOx (2µmol mol-1 NO and c. 0.5 µmol mol-1 NO2 ) to CO2 -enriched air significantly reduced the yield, compared with the 'clean, high CO2 treatment, without producing visible symptoms of toxicity. Fumigation of single plants in high CO2 , with NOx rapidly inhibited photosynthesis per unit leaf area. This did not appear to be due to a reduction in stomatal conductance. Removal of NOx from the atmosphere caused a rapid and complete recovery in the rate of photosynthesis. Studies were made of the effects of growing Plants for long periods in atmospheres containing high CO2 and NOx on the photosynthetic capacity of single leaves when measured in NOx -free air. The decrease in photosynthetic rate as the fourth leaf aged occurred earlier in plants grown in CO2 -enriched air than in those from the low CO2 treatment. Leaves which developed in the CO2 -enriched air containing NOx did not suffer any long-term damage to photosynthetic activity in comparison with those of the 'clean' high CO2 . In mature leaves the principal long-term effect of enrichment (with or without NOx was to reduce the rate of photosynthesis in saturating CO2 . In contrast, there was less effect on the rate of Photosynthesis in low CO2 . The absence of a long-term effect of NOx , on the photosynthetic capacity suggested that photosynthesis by the lettuce crop will be inhibited only during the transient periods of NOx accumulation in the glasshouse.