Slow recovery of High Arctic heath communities from nitrogen enrichment

Acknowledgements We are indebted to Ian Alexander who initiated the fertilisation experiment with SJW; to successive researchers involved in the experiment, John Baddeley, Nanette Madan, Lars Hogbom, Bernard Moyersen, Carmen Gordon; and to field assistants, Alison Horsburgh, Andrew Coughlan, Jo Wynn, Lora Crabtree. We thank Hans Kruijer and Michael Stech for assistance with bryophyte species identification in 2011. Funding for the initial experiment was provided by the NERC Arctic Terrestrial Ecology Special Topic Programme (GR3/9424, GR9/3433) with additional support from the CEC TMR Programme, Ny-Ålesund LSF and the British Ecological Society. This recovery study was funded by NERC (NE/I016899/1). The research was made possible by use of NERC facilities at Harland Huset; special thanks to Nick Cox and colleagues for their unfailing hospitality and support.Peer reviewedPostprintPostprin

Successional Change in Phosphorus Stoichiometry Explains the Inverse Relationship between Herbivory and Lupin Density on Mount St. Helens

The average nitrogen-to-phosphorus ratio (N?P) of insect herbivores is less than that of leaves, suggesting that P may mediate plant-insect interactions more often than appreciated. We investigated whether succession-related heterogeneity in N and P stoichiometry influences herbivore performance on N-fixing lupin (Lupinus lepidus) colonizing primary successional volcanic surfaces, where the abundances of several specialist lepidopteran herbivores are inversely related to lupin density and are known to alter lupin colonization dynamics. We examined larval performance in response to leaf nutritional characteristics using gelechiid and pyralid leaf-tiers, and a noctuid leaf-cutter.Apple JL, Wink M, Wills SE, Bishop JG (2009) Successional Change in Phosphorus Stoichiometry Explains the Inverse Relationship between Herbivory and Lupin Density on Mount St. Helens. PLoS ONE 4(11): e7807. doi:10.1371/journal.pone.000780

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

Above-and belowground insect herbivory modifies the response of a grassland plant community to nitrogen eutrophication

Understanding the role that species interactions play in determining the rate and direction of ecosystem change due to nitrogen (N) eutrophication is important for predicting the consequences of global change. Insects might play a major role in this context. They consume substantial amounts of plant biomass and can alter competitive interactions among plants, indirectly shaping plant community composition. Nitrogen eutrophication affects plant communities globally, but there is limited experimental evidence of how insect herbivory modifies plant community response to raised N levels. Even less is known about the roles of above‐ and belowground herbivory in shaping plant communities, and how the interaction between the two might modify a plant community's response to N eutrophication. We conducted a 3‐yr field experiment where grassland plant communities were subjected to above‐ and belowground insect herbivory with and without N addition, in a full‐factorial design. We found that herbivory modified plant community responses to N addition. Aboveground herbivory decreased aboveground plant community biomass by 21%, but only at elevated N. When combined, above‐ and belowground herbivory had a stronger negative effect on plant community biomass at ambient N (11% decrease) than at elevated N (4% decrease). In addition, herbivory shifted the functional composition of the plant community, and the magnitude of the shifts depended on the N level. The N and herbivory treatments synergistically conferred a competitive advantage to forbs, which benefited when both herbivory types were present at elevated N. Evenness among the plant species groups increased when aboveground herbivory was present, but N addition attenuated this increase. Our results demonstrate that a deeper understanding of how plant–herbivore interactions above and below ground shape the composition of a plant community is crucial for making reliable predictions about the ecological consequences of global change

Photosynthetic performance in Sphagnum transplanted along a latitudinal nitrogen deposition gradient

Increased N deposition in Europe has affected mire ecosystems. However, knowledge on the physiological responses is poor. We measured photosynthetic responses to increasing N deposition in two peatmoss species (Sphagnum balticum and Sphagnum fuscum) from a 3-year, north-south transplant experiment in northern Europe, covering a latitudinal N deposition gradient ranging from 0.28 g N m(-2) year(-1) in the north, to 1.49 g N m(-2) year(-1) in the south. The maximum photosynthetic rate (NPmax) increased southwards, and was mainly explained by tissue N concentration, secondly by allocation of N to the photosynthesis, and to a lesser degree by modified photosystem II activity (variable fluorescence/maximum fluorescence yield). Although climatic factors may have contributed, these results were most likely attributable to an increase in N deposition southwards. For S. fuscum, photosynthetic rate continued to increase up to a deposition level of 1.49 g N m(-2) year(-1), but for S. balticum it seemed to level out at 1.14 g N m(-2) year(-1). The results for S. balticum suggested that transplants from different origin (with low or intermediate N deposition) respond differently to high N deposition. This indicates that Sphagnum species may be able to adapt or physiologically adjust to high N deposition. Our results also suggest that S. balticum might be more sensitive to N deposition than S. fuscum. Surprisingly, NPmax was not (S. balticum), or only weakly (S. fuscum) correlated with biomass production, indicating that production is to a great extent is governed by factors other than the photosynthetic capacity

New science on the effects of nitrogen deposition and concentrations on Natura 2000 sites (theme 3): background documement

This background paper summarizes established and new science on the effects of nitrogen (N) deposition on ecosystems and considers the potential for improved assessment of N deposition impacts on Natura 2000 sites. The key aspects covered are N deposition effects on biodiversity and on biogeochemistry, links to ecosystem services, the importance of N form, N deposition indicators, management practices and ecosystem reversibility following decreased N input. The paper shows that: • Evidence of N impacts on different vegetation types in Europe exists, but that it is important that it is translated meaningfully to the target habitats listed under the Habitats Directive. Evidence for N deposition effects to important ecosystem services, such as carbon sequestration, also exists but the cause and effect relationships underlying them are often complex and not sufficiently understood. • Chemical N form can influence both the rate of ecosystem change, and the extent of impacts on the short and long-term. Evidence is presented for ammonia causing detrimental plant physiological effects, probably on the majority of species, whilst ammonium and nitrate effects will depend on plant species present. • Plant species composition as well as plant biochemical parameters may be useful bioindicators for assessment of N deposition effects in Natura 2000 sites, however “baseline” data are mostly not available for rare species. Ecosystem specific indicators, that are predictive of further damage, rather than a consequence of already existing adverse effects (i.e., early warning indicators) are needed. • Site level management practices can be useful to reduce the impact if N deposition but they will certainly not be able to mitigate all the impacts of enhanced N deposition and enhances N concentration on Natura 2000 habitats. More knowledge is needed to better understand where and if management intervention is appropriate to mitigate N effects. • Studies on the reversibility of N impacts show that some ecosystem parameters may revert quickly, while other components may show strong inertia. In some cases reversion to the original state may however be impossible. • Climatic factors interfere with ecosystem effects of N deposition. It is clear that climate both can emphasize and mitigate effects of N deposition. Current climate and expected climatic changes must be included in assessments and predictions of N deposition effects on ecosystems. The aim is to provide a broad picture of scientific advancement within the field of N deposition research, and to provide a starting point for workshop discussions. Workshop discussions addressed the relevance of new science in assessing N deposition impacts on Natura 2000 sites, and identified when there is potential to make management adjustments to mitigate such effects

The impact of wildfire on biogeochemical fluxes and water quality in boreal catchments

Wildfires are the major disturbance in boreal ecosystems and are of great importance for the biogeochemical cycles of carbon (C) and nutrients. However, these fire-induced impacts are hard to quantify and are rarely assessed together at an ecosystem level incorporating both aquatic and terrestrial environments. Following a wildfire in Sweden in an area with ongoing monitoring, we conducted a pre-fire (9 years) and post-fire (4 years) multi-catchment investigation of element losses (combustion and leaching) and impacts on water quality. Direct C and nitrogen (N) losses through combustion were ca. 4500 and 100 g m−2, respectively. Net CO2 loss associated with soil and biomass respiration was ∼ 150 g C m−2 during the first year, but the ecosystem started to show net CO2 uptake in June 3 years post-fire. Aquatic C and N losses the first 12 months post-fire were 7 and 0.6 g m−2, respectively. Hence, soil respiration comprised a non-negligible part of the post-fire C loss, whereas aquatic C losses were minor and did not increase post-fire. However, other elements (e.g. Ca, S) exhibited ecologically relevant increases in fluvial export and concentration with large peaks in the immediate post-fire period. The temporal dynamics of stream concentrations (Ca2+, Mg2+, K+ ,SO−24, Cl− ,NH+4, total organic N) suggest the presence of faster- and slower-release nutrient pools with half-lives of around 2 weeks and 4 months which we attribute to physicochemically and biologically mediated mobilization processes, respectively. Three years after the fire, it appears that dissolved fluxes of nutrients have largely returned to pre-fire conditions, but there is still net release of CO2