Montane heathland rejuvenation by choppering—Effects on vascular plant and arthropod assemblages

Land-use changes and atmospheric nitrogen deposition have negatively affected heathland biota. Active habitat management is one possible way of counteracting the biodiversity loss associated with these habitat alterations. However, management practices for lowland heathlands often have been transferred to montane heathlands, irrespective of the differences in environmental conditions or assemblage composition. The objective of this study was to evaluate the effects of so-called choppering for the rejuvenation of montane heathland. Choppering involves chaffing and removing the largest part of the organic layer down to the mineral soil. In this study, we compared montane heathlands that were rejuvenated through the application of choppering (CHOPPER) to old-growth montane heathlands (CONTROL). Thirteen years after the rejuvenation measures had been conducted, the environmental conditions between CHOPPER and CONTROL still differed. CHOPPER was characterised by shorter vegetation (herbs/grasses and dwarf shrubs), more bare soil, less litter and higher temperatures. Although, the vascular plants and all studied arthropod groups were affected by the environmental changes, their responses were somewhat different. CHOPPER had a unique assemblage of each taxonomic group that included at least a few heathland species that mainly occurred in this treatment. However, choppering was most beneficial for vascular plants, grasshoppers and carabid beetles. As shown for lowland heathlands, choppering is also a suitable management measure for montane heathland to rejuvenate vegetation with its characteristic arthropod fauna. Due to the intact seed banks and Ericaceae root systems with their mycorrhizas, in combination with the availability of bare soil, heathland vegetation can rapidly regenerate after choppering. The keystone structures that explain the high relevance of CHOPPER, especially for vascular plants, grasshoppers and carabid beetles, are low-growing vegetation and bare soil, which result in light and warm microclimatic conditions. Based on the results of our study, we recommend choppering as a regular management measure to rejuvenate montane heathland.     

Waiting for fungi: the ectomycorrhizal invasion of lowland heathlands

1.  In England, the loss of lowland heathland, a habitat of global conservation importance, is primarily due to the invasion of birch and pine. This encroachment has been researched in depth from a plant perspective but little is known about the role of mycorrhizal fungi. In lowland heathlands the resident dwarf shrubs form ericoid mycorrhizas whereas invading trees form ectomycorrhizas. Therefore, tree encroachment into heathlands can be regarded as the replacement of a resident mycorrhizal community by an invading one.
2 . This study examined how fungi form mycorrhizas with Betula and Pinus in lowland heathlands. We addressed the question of whether there are mycorrhizal fungi that mediate invasion using a molecular ecology approach to compare the mycorrhizal inoculum potential of soil at three levels of invasion (uninvaded heathland, invaded heathland and woodland) and the fungi forming mycorrhizas on tree seedlings and trees across diverse sites.
3.  We show that in lowland heathlands: (i) seedlings have severely limited access to ectomycorrhizal fungi relative to woodlands, (ii) there are few keystone spore-dispersed ectomycorrhizal fungi that can mediate tree invasion, (iii) tree seedlings can remain non-mycorrhizal for at least one year when no inoculum is present, even near saplings, and (iv) mycorrhizal seedlings achieve greater biomass than non-mycorrhizal seedlings. Within uninvaded heathland we detected only Rhizopogon luteolus , Suillus variegatus , S. bovinus ( Pinus symbionts) and Laccaria proxima (primarily a Betula symbiont).
4. Synthesis . Overall, ectomycorrhizal inoculum in lowland heathlands is rare; most tree seedlings growing in heathland soil are not mycorrhizal due to limited spore dispersal, poorly developed spore banks and weak common mycorrhizal networks. These seedlings can persist awaiting mycorrhization to boost their growth.     

Spider communities as evaluation tools for wet heathland restoration

Over the last two centuries wet heathlands and associated habitats, such as poor fens and bogs, have suffered extensive fragmentation. Recently, large-scale projects aim to restore these rare habitats throughout Europe. To evaluate post-restoration trajectories of wet heathlands, suitable monitoring tools are urgently needed. Here, we investigated whether spider communities are useful tools for evaluating the restoration success of wet heathlands. Ordination revealed that dissimilarity in spider communities between patches of different age classes resulted mainly from vegetation cover and soil moisture. By using a functional trait-approach, we tested if the time since restoration affects trait distribution of spiders in wet heathlands. Typical wet heathland spider species were less common with increasing vegetation encroachment and lower water content. New patches were inhabited by summer active, eurytopic (non-heathland) spiders, while more typical heathland species were found in middle-aged and old patches. Our results suggest that time-related changes in vegetation structure and moistness of restored wet heathlands are clearly reflected by spider communities. Although mobile spiders quickly recolonize the restored heathlands, it takes time for typical heathland spiders to settle. Restoration measures should prevent the negative effects of a vegetation encroachment and a high density of forested edges and should rehabilitate the hydrological cycle in order to preserve rare heathland spiders. We discuss that accounting for responses of spiders provides additional information to guide wet heathlands restoration.     

Fire increases aboveground biomass, seed production and recruitment success of Molinia caerulea in dry heathland

During the last decades, the perennial tussock grass Molinia caerulea has shown an increased abundance in European heathlands, most likely as a result of increased nitrogen deposition and altered management schemes. Because of its deciduous nature, Molinia produces large amounts of litter each year, which may affect the intensity and frequency of accidental fires in heathlands. These fires may influence plant population dynamics and heathland community organization through their effects on plant vital attributes and competitive interactions. In this study, fire-induced changes in competitive ability and invasiveness of Molinia through changes in biomass production, seed set and seed germination under both natural and laboratory conditions were investigated. We found that fire significantly increased aboveground biomass, seed set and germination of Molinia. Seed set was twice as high in burned compared to unburned heathland. Two years after fire, seedling densities in natural conditions were on average six times higher in burned than in unburned heathland, which resulted in increased abundance of Molinia after burning. The seed germination experiment indicated that seeds harvested from plants in burned heathland showed higher germination rates than those from unburned heathland. Hence, our results clearly demonstrate increased invasive spread of Molinia after large and intense fires. Active management guidelines are required to prevent further encroachment of Molinia and to lower the probability of large fires altering the heathland community in the future.     

Effects of Montane Heathland Restoration on Leafhopper Assemblages (Insecta: Auchenorrhyncha)

At the beginning of the 20th century, many montane heathlands were abandoned and became subject to natural succession or afforestation by humans. Thus, the formerly large montane heathlands slowly degraded into small and isolated patches. In this study, we evaluate the influence of restoration measures on leafhopper (Auchenorrhyncha) assemblages of montane heathland ecosystems in Central Europe. Our analyses comprised three different site types that were adjacent to each other: (1) montane heathlands, (2) restoration sites, and (3) control sites. Leafhoppers showed a clear response to montane heathland restoration. Thus, after 4–5 years since implementation of restoration measurements restoration sites were characterized by the highest species richness. However, detailed analyses of leafhopper diversity, species composition, and environmental parameters on the three site types revealed that restoration sites were rather similar to control sites and significantly differing from montane heathlands. We conclude that leafhoppers are excellent bioindicators for restoration measurements because they reflected environmental differences between the three site types. Restoration measurements might only be a useful instrument to promote typical montane heathland leafhopper communities in the long run. Colonization by leafhoppers is, however, dependent on many different factors such as leafhopper mobility, vegetation structure, microclimate, and the establishment of ericaceous dwarf shrubs. Practitioners should establish a management regime (grazing and sod‐cutting) that creates a mosaic of different habitat structures and increases typical heathland vegetation, thus, favoring the colonization of typical heathland leafhoppers.     

Genetic erosion in a stenotopic heathland ground beetle (Coleoptera: Carabidae): a matter of habitat size?

Two centuries ago large areas of north-west Europe were covered by coherent heathlands which hosted numerous specialized species. Changes in land use made heathlands fragmented and rare, consequently, they are in the focus of nature conservation efforts today. But how large should remaining heathland patches be in order to secure the survival of populations of specialized species? We investigated the genetic diversity at five allozyme loci of Poecilus lepidus, a flightless and stenotopic heathland ground beetle. 29 populations from differently sized heathland patches in north-west Germany were analyzed. Results show a weak but significant genetic differentiation and no evidence for isolation by distance or other patterns of spatial autocorrelation. Linear regression analysis revealed significant relationships between patch size, allelic richness, number of alleles and expected heterozygosity. These findings are explained by severe habitat fragmentation together with strong fluctuations in population size which have been reported for this species in the past. To conserve the vast majority of the species’ genetic diversity for a period of 100 years we suggest to maintain heathland patches of at least 50 ha in size.     

Fire increases aboveground biomass,seed production and recruitment success of Molinia caerulea in dry heathland

During the last decades, the perennial tussock grass Molinia caerulea has shown an increased abundance in European heathlands, most likely as a result of increased nitrogen deposition and altered management schemes. Because of its deciduous nature, Molinia produces large amounts of litter each year, which may affect the intensity and frequency of accidental fires in heathlands. These fires may influence plant population dynamics and heathland community organization through their effects on plant vital attributes and competitive interactions. In this study, fire-induced changes in competitive ability and invasiveness of Molinia through changes in biomass production, seed set and seed germination under both natural and laboratory conditions were investigated. We found that fire significantly increased aboveground biomass, seed set and germination of Molinia. Seed set was twice as high in burned compared to unburned heathland. Two years after fire, seedling densities in natural conditions were on average six times higher in burned than in unburned heathland, which resulted in increased abundance of Molinia after burning. The seed germination experiment indicated that seeds harvested from plants in burned heathland showed higher germination rates than those from unburned heathland. Hence, our results clearly demonstrate increased invasive spread of Molinia after large and intense fires. Active management guidelines are required to prevent further encroachment of Molinia and to lower the probability of large fires altering the heathland community in the future.     

The effect of topsoil removal in restored heathland on soil fauna, topsoil microstructure, and cellulose decomposition: implications for ecosystem restoration

Communities of soil macrofauna, oribatid mites, and nematodes as well as vegetation and soil chemistry were studied on twelve plots representing three replicates of the following treatments: agricultural meadow, heathland, and heathland restored either by partial or complete topsoil removal 15 years earlier. We also studied the effect of soil macrofauna on decomposition and the microstructure of the soil surface layer with litterbags in combination with the analysis of thin soil sections. The communities of soil macrofauna and oribatid mites significantly differed between agricultural meadows and heathlands. The partial and complete topsoil removal plots were more similar to heathlands with respect to macrofauna and to agricultural meadows with respect to oribatid mites. The density and diversity of soil macrofauna was higher in agricultural meadows than in heathlands; in particular, earthworms, litter transformers, root feeders, and microsaprophags were more abundant on meadows. Heathlands, in contrast, contained a much higher diversity of oribatid mites. The community structure of nematodes did not significantly differ among the treatments. Analysis of thin soil sections revealed a thick organic fermentation layer in heathlands, which was absent in agricultural meadows and only weakly developed in the topsoil removal plots. In agricultural meadows, litterbags and thin soil sections indicated that abundant macrofauna, including endogeic earthworms, were very effective in removing the litter from the soil surface and mixing it into the mineral soil. Possible effects of this soil mixing on restoration success are discussed.     

Salt spray differentially affects water status, necrosis, and growth in coastal sandplain heathland species

Sandplain heathlands are disturbance-dependent plant communities that occur infrequently in coastal areas of the northeastern United States. We hypothesize that salt spray plays a role in maintaining the composition of the heathland community by excluding salt-intolerant species close to the ocean. We examined the distributions of Solidago nemoralis, Myrica pensylvanica, Pinus rigida, and Quercus spp. in heathlands and conducted greenhouse studies to determine whether different levels of salt spray tolerance explain patterns found in the field. We found that common heathland forb and shrub species grow closer to the ocean than successional woody species. In greenhouse experiments, these species differ in their water status, necrosis, and growth responses to salt spray. The tree species P. rigida and Q. rubra are more susceptible to salt spray than the common heathland species M. pensylvanica. Our results suggest that salt spray may prevent tree species in heathlands from growing close to the ocean and therefore might be an important factor in maintaining the characteristic community composition of these dwarf shrublands in coastal habitats.     

Impacts of succession and grass encroachment on heathland Orthoptera

For the conservation of biodiversity, heathlands present important ecosystems throughout Europe. The formerly widespread habitats are nowadays restricted to small and isolated remnants. Without land use heathland vegetation undergoes succession and, in addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses. In the present study we analysed the effects of succession and grass encroachment on Orthoptera in a coastal heathland on the Baltic island of Hiddensee, Germany. Vegetation, microclimate, soil humidity and Orthoptera were sampled in the five main stages of heathland succession, namely grey dunes, dwarf-shrub heath, grassy heath, heath with shrubs, and birch forest. Vegetation and environmental parameters showed strong differences among the successional stages. Orthoptera species richness was highest in transitional stages. The high proportion of grasses offer favourable habitat conditions for graminivorous, chorto- and thamnobiont species. Orthoptera density was highest in grey dunes. Threatened and specialised species were restricted to the young stages grey dunes and dwarf-shrub heath. Hence, in order to maintain a high diversity of Orthoptera in heathlands, maintaining different successional stages is of critical importance and this should be integrated into heathland management practices.     

Salt spray and edaphic factors maintain dwarf stature and community composition in coastal sandplain heathlands

North American coastal sandplain heathlands are unique in species composition and vegetation, but the extent to which edaphic factors influence the structure of these communities is currently debated. It was hypothesized that salt spray and edaphic factors maintain the dwarf stature and community composition of heathlands by limiting plant growth and excluding competitively dominant woody species close to the ocean. Field surveys were carried out to investigate the spatial patterns of salt spray accumulation, soil salt and soil moisture. High salt spray correlated significantly with increased leaf necrosis and water stress in Myrica pensylvanica and with decreased plant height. Plant community composition changed across a salt spray and soil gradient, as well. Distinctive sub-communities were identified that separated according to soil salt and soil moisture but salt spray was the main factor affecting sites occupied only by heathland vegetation. Results from this study suggest that salt spray suppresses the growth of heathland plants in close proximity to the ocean, and therefore maintains the low stature in these dwarf shrublands. This research also demonstrates that the physical environment influences the community structure in heathlands, particularly by limiting tree species from growing in high salt spray, low water availability sites.     

Coastal heathland vegetation is surprisingly resistant to experimental drought across successional stages and latitude

In the last decade, several major dwarf-shrub dieback events have occurred in northern European coastal heathlands. These dieback events occur after extended periods with sub-zero temperatures under snow-free conditions and clear skies, suggesting that coastal heathlands have low resistance to winter drought. As climate projections forecast increased drought frequency, intensity, and duration, coastal heathlands are likely to experience more such diebacks in the future. There are, however, few empirical studies of drought impacts and responses on plant communities in humid oceanic ecosystems. We established a drought experiment with two distinct levels of intensified drought to identify responses and thresholds of drought resistance in coastal heathland vegetation. We repeated the experiment in two regions, separated by five degrees latitude, to represent different bioclimatic conditions within the coastal heathlands' wide latitudinal range in Europe. As coastal heathlands are semi-natural habitats managed by prescribed fire, and we repeated the experiment across three post-fire successional phases within each region. Plant community structure, annual primary production, and primary and secondary growth of the dominant dwarf-shrub Calluna vulgaris varied between climate regions. To our surprise, these wide-ranging vegetation- and plant-level response variables were largely unaffected by the drought treatments. Consequently, our results suggest that northern, coastal heathland vegetation is relatively resistant to substantial intensification in drought. This experiment represents the world's wettest (2200 mm year⁻¹) and northernmost (65°8'N) drought experiment to date, thus filling important knowledge gaps on ecological drought responses in high-precipitation and high-latitude ecosystems across multiple phases of plant community succession.     

Heather and heathlands

GIMINGHAM, C. H., 1989. Heather and heathlands. Studies of the biology and ecology of heather (Calluna vulgaris (L.) Hull) reveal a remarkable combination of characteristics, accounting for its success as a heathland dominant and its ability to persist under traditional forms of use and management. New work on the life history and physiology of the species is helping to explain recent changes in heathlands, and to develop appropriate methods of conservation.     

Catchment Liming to Restore Degraded, Acidified Heathlands and Moorland Pools

Current restoration measures of degraded, acidified heathland ecosystems have not always been successful in the Netherlands. Positive effects of a restored hydrology are often counteracted by acidification of the soil and the local groundwater system. Liming of the heathlands in the catchment of moorland pools might contribute to the restoration of both habitats. Experimental catchment liming was carried out in two degraded Dutch heathlands, with doses varying between 2 and 6 tons/ha. Catchment liming resulted in increased pH and base cation concentrations in the highest elevated limed parts, as well as in the lower situated, nonlimed heath areas and moorland pools. Generally, catchment liming created suitable conditions for the return of heathland target species, and the positive effects lasted for at least 6 years. The response of the heathland vegetation to the liming has, however, been slow because only a small number of endangered plant species increased in abundance. In contrast, four Red List soft‐water macrophytes strongly increased in abundance in the moorland pool. Our results show that, even with the slow return of Red List plant species, catchment liming can be a successful management tool for the restoration of the acidified heathland landscape.     

Rapid response of Orthoptera to restoration of montane heathland

Understanding how to restore threatened ecosystems is of special relevance for nature conservation. The aim of this study was to use Orthoptera as ecological indicators for the effects of montane heathland restoration in Central Europe. We analysed the three following treatments: (i) montane heathlands (MONHEATH) (N = 7), (ii) restoration sites (RESSITE) (N = 3) and (iii) clear-cuts of spruce forests as unprocessed and ungrazed control sites (CONTROL) (N = 3). Vegetation structure and microclimate differed considerably between MONHEATH on the one hand and RESSITE and CONTROL on the other hand. Orthoptera species richness and density did so too. MONHEATH was characterised by a high-growing dense dwarf-shrub and moss layer having a cool microclimate and high soil moisture. In contrast, RESSITE and CONTROL had sparse vegetation and a warm microclimate; Orthoptera species richness and density was highest on these sites. Our study clearly showed that heathland Orthoptera responded rapidly to restoration measures, while Ericaceae dwarf shrubs slowly established. The vast majority of Orthoptera species found on the restoration sites are early and mid-successional species. The colonization of the sites by late-successional Orthoptera species in the future will depend on the further development of the heathland vegetation; that is, if Ericaceae will expand to the sites. We conclude that the realised restoration measures are suitable to promote heathland Orthoptera of early and mid-successional stages. However, the current management of montane heathlands is insufficient and needs to be intensified in order to provide structurally diverse habitats with their characteristic orthopteran assemblages.     

Effects of afforestation with pines on woody plant diversity of Mediterranean heathlands in southern Spain

Open heathlands of the Strait of Gibraltar region constitute a singularvegetation type within the Mediterranean Basin owing to their high biodiversitylevels and distinct features in the biological aspects of endemism. However,they have been traditionally depicted as low profitable, tree-less, barrenlandsand have been either overlooked in conservation policies or included inextensive pine-tree afforestation programmes. Nowadays, most of the existingpine plantations are no longer exploited and have become dense pine woodlands.Here we present the results of a comparative analysis aimed to ascertain theeffects of pine stands on the structure and diversity of the woody component ofthese mediterranean heathlands. Ten sampling sites were located within thestudyarea, each consisting of two adjacent subsamples: an open heathland and aneighbouring heathland under pine-tree cover. Woody species richness andoverallshrub cover was much lower in pine-tree heath understoreys. This drop in coverwas more marked for short-lived species (seeders and withering resprouters),implying a decrease of the life-history diversity and also having noticeableconsequences in homogenizing the floristic composition of heath patches underpine trees. Heathland narrow endemics dropped significantly in pine stands,showing greater sensitivity to afforestation than nonendemics. We suggest thatthe open heathlands of the Strait of Gibraltar should be given furtherattentionin EU and IUCN conservation policies. Programmes should be implemented toprogressively eliminate abandoned pine plantations so as to restore andpreservethese highly diverse mediterranean heathlands.     

Conservation of heathland by sheep grazing in Brittany (France): Importance of grazing period on dry and mesophilous heathlands

Heathlands are characteristic semi-natural ecosystems of western Europe. Nowadays they are threatened because of their lack of agricultural value. Management of the remaining heathland areas necessitates precise knowledge about different management tools that can be used, including grazing. The effects of sheep grazing on dry and mesophilous heathlands of Brittany were studied in relation to grazing season and duration. Management aims were to control shrubs and to maintain the dominance of heathers. The grazing pressures applied here were quite high, about 10–15 sheep ha⁻¹ yr⁻¹. Erica ciliaris appeared to be very sensitive to grazing at the end of summer. This implicates special care in the management of mesophilous heathland. In dry heathland, Erica cinerea did not show this sensitivity. Control of shrubs was achieved by most grazing regimes which led to a decrease of 11–22 cm in gorse height, except summer grazing which seemed to stimulate their growth.This study underlined the relevance of the use of sheep grazing as a heathland management tool but also the necessity of prior experiments to adapt these operations mainly in terms of grazing period to the type of heathland and the objectives put forward.     

Responses of common and successional heathland species to manipulated salt spray and water availability

Coastal sandplain heathlands are a rare plant community in the northeastern United States. Salt spray and water availability are likely important factors determining heathland distribution. Field surveys and manipulative experiments were performed to examine heathland species' responses to salt spray and water availability. We surveyed field distributions of four typical heathland species: Solidago puberula, Solidago rugosa, Gaylussacia baccata, and Myrica pensylvanica. The distributions of two native tree species, Pinus rigida and Quercus ilicifolia, were also surveyed because they succeed into coastal heathlands with low disturbance frequency. We then manipulated salt spray and water in the field and measured species' water status, necrosis, and growth responses to the treatments. Predawn xylem pressure potential and necrosis were strongly affected by high salt spray and low water availability. Shoot elongation was also limited in S. puberula and S. rugosa grown in high salt, low water treatments. Gaylussacia baccata and Q. ilicifolia were particularly sensitive to high salt spray and low water, suggesting that they might excluded be from areas with those conditions. The interaction between salt spray and water availability could affect the landscape scale and should be incorporated into conservation management plans.     

Management and diversity of arthropods inCalluna heathland

Heathlands, dominated byCalluna vulgaris, are located along the western seaboard of Europe. Most undergo a cyclical succession as the heather plants pass through four definable growth-phases over a period of about 30 years. Examples of upland heathlands are drawn from the North York Moors National Park, an area of Jurassic rocks in the north of England (approximately 54°25N, 0°55W). Botanical diversity is low. In a survey of 10 heathlands selected to encompass the botanical variation of the heathlands of the National Park, only 40 species of higher plants and 31 species of mosses were recorded (2.2% and 4.5% of the British flora, respectively).However, diversity of some invertebrate groups is high. Totals of 54 species of carabid beetles and 127 species of spiders were recorded (15.3% and 20.4% of the British fauna, respectively). At least two environmental gradients appear to influence the species composition of the invertebrate assemblages: a wet-dry gradient and the height of the vegetation (corresponding to the growth-phase of the heather). Experimental areas on three heathlands were burnt and cut, the traditional and more modern methods of heather management. Analysis of both the spider and beetle data indicated that the species assemblages were primarily influenced by the growth-phase of the heather. Some of the nationally rarer species are associated with the open conditions of recently-cut or burnt heathland. An altitudinal effect was also inferred since heathland at 410 m above sea level tended to be less species-rich than heathlands at ca 260 m above sea level.These results present a paradox. Although botanical diversity is extremely low, and although it is a non-natural ecosystem maintained by fire and grazing management to prevent a succession to woodland, arthropod diversity can be extremely high. The discussion focuses on the importance of upland heathlands in the conservation of insect and spider diversity, as well as on the value of these upland areas for conservation more generally.C. vulgaris is seen to be a key species on which many other species depend. One conclusion is that there is insufficient effort, nationally and especially internationally, to conserve the heathlands of Europe.     

Effect of plant succession, ploughing, and fertilization on the microbiological oxidation of atmospheric methane in a heathland soil

Abstract: The effect of plant succession on methane uptake was measured on intact soil cores collected from seven heathland sites. Six of the sites had undergone either secondary succession with grass or oak, ammonium fertilization or ploughing, while the seventh site was located in the native heathland. There was a positive relationship between methane uptake rate and time elapsed since the plant invasion had taken place in the native heathland. The native heathland site showed an insignificant atmospheric methane uptake of 0.01 mg CH₄ m⁻² d⁻¹, whereas the established oak brushwood (70 years old) and the grass invaded heathland (13 years old) showed rates of 1.36 mg CH₄ m⁻² d⁻¹ and 0.73 mg CH₄ m⁻² d⁻¹, respectively. In the fertilized heathland plot (112 kg N ha⁻¹ six years prior to this study) grass had become the dominating species and showed a methane oxidation rate of 0.28 mg CH₄ m⁻² d⁻¹. Ploughing of the heathland resulted in methane oxidation rates seven times the rates measured in the native heathland. The results suggested that an increased future atmospheric nitrogen deposition in heathlands and other nutrient poor ecosystems may have a stimulating effect on the soil sink for atmospheric methane.