Andrea J. Britton

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.

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.

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.

Terricolous alpine lichens are sensitive to both load and concentration of applied nitrogen and have potential as bioindicators of nitrogen deposition.

The influence of applied nitrogen (N) concentration and load on thallus chemistry and growth of five terricolous alpine lichen species was investigated in a three-month N addition study. Thallus N content was influenced by both concentration and load; but the relative importance of these parameters varied between species. Growth was most affected by concentration. Thresholds for effects observed in this study support a low critical load for terricolous lichen communities (<7.5 kg N ha(-1) y(-1)) and suggest that concentrations of N currently encountered in UK cloudwater may have detrimental effects on the growth of sensitive species. The significance of N concentration effects on sensitive species also highlights the need to avoid artificially high concentrations when designing N addition experiments. Given the sensitivity of some species to extremely low loads and concentrations of N we suggest that terricolous lichens have potential as indicators of deposition and impact in northern and alpine ecosystems.

Drought alters carbon fluxes in alpine snowbed ecosystems through contrasting impacts on graminoids and forbs

• Climate change is predicted to increase the frequency of drought events in alpine ecosystems with the potential to affect carbon turnover. • We removed intact turfs from a Nardus stricta alpine snowbed community and subjected half of them to two drought events of 8 d duration under controlled conditions. Leachate dissolved organic carbon (DOC) was measured throughout the 6 wk study period, and a (13)CO(2) pulse enabled quantification of fluxes of recent assimilate into shoots, roots and leachate and ecosystem CO(2) exchange. • The amount of DOC in leachate from droughted cores was 62% less than in controls. Drought reduced graminoid biomass, increased forb biomass, had no effect on bryophytes, and led to an overall decrease in total above-ground biomass compared with controls. Net CO(2) exchange, gross photosynthesis and the amount of (13)CO(2) fixed were all significantly less in droughted turfs. These turfs also retained proportionally more (13)C in shoots, allocated less (13)C to roots, and the amount of dissolved organic (13)C recovered in leachate was 57% less than in controls. • Our data show that drought events can have significant impacts on ecosystem carbon fluxes, and that the principal mechanism behind this is probably changes in the relative abundance of forbs and grasses.

Additive impacts of nitrogen deposition and grazing on a mountain moss-sedge heath

High levels of nitrogen (N) deposition and long-term heavy grazing pressures are thought to have caused severe degradation and loss of alpine habitats in the UK. Reductions in stocking density following agricultural reform provide an opportunity for restoration but it is unclear to what extent this may be limited by continuing N deposition. We investigated the effects of spatial variability in grazing pressure on the tissue chemistry and growth of the key bryophyte Racomitrium lanuginosum at a moss-heath site in north Wales exposed to high N deposition. Racomitrium N content and cell membrane damage both clearly reflected the high atmospheric deposition and the additional effect of N deposited by grazers, but this was not translated into negative effects of either factor on growth. Racomitrium grew fastest where a well-developed moss mat was present, regardless of grazer presence, and grew slower in degraded areas where the mat was absent or fragmentary. Absence of a moss mat may produce conditions where Racomitrium is more frequently desiccated, resulting in low growth rates and perpetuating the poor condition of the moss heath. Removal of grazing may result in an improvement in Racomitrium condition, even under existing levels of N deposition, where the moss mat remains, but restoration of the most damaged areas may require further intervention.ZusammenfassungAtmosphärischer Stickstoffeintrag (N Deposition) und intensive Beweidung haben zu Beeinträchtigung und Verlust der alpinen Habitate in Grossbritannien geführt. Eine Verminderung der Bestockungsdichten infolge einer Landwirtschaftsreform könnte die Regeneration solcher Habitate begünstigen, doch ob eine Regeneration bei bleibend hoher N Deposition überhaupt möglich ist, ist unklar. In einem alpinen Heidegebiet in Nord-Wales mit hohen N Depositionsraten untersuchten wir, wie sich verschiedene Beweidungsintensitäten auf das Wachstum und die chemische Zusammensetzung des Mooses Racomitrium lanuginosum, der dominierenden Art dieser Pflanzengesellschaft, auswirken. Der N Gehalt und die Zellmembran-Integrität von Racomitrium widerspiegelten klar die hohe atmosphärische N Deposition und den zusätzlichen N Eintrag durch Beweidung, aber keiner dieser Faktoren führte zu vermindertem Wachstum. Unabhängig von der Beweidungsintensität wuchs Racomitrium da am besten, wo eine dichte Moosbedeckung vorhanden war, und schlechter in gestörten Standorten mit fehlender oder fragmentierter Moosdecke. Möglicherweise trocknet Racomitrium an solch gestörten Standorten häufiger aus, was das Wachstum bremst und die Regeneration der Moosdecke verlangsamt. Eine Aufgabe der Beweidung kann also auch bei anhaltend hoher N Deposition den Zustand von Racomitrium verbessern, wenn noch eine intakte Moosbedeckung vorhanden ist. Um bereits geschädigte Racomitrium-Heide zu regenerieren, sind jedoch zusätzliche Massnahmen nötig.

Controls on soil solution nitrogen along an altitudinal gradient in the Scottish uplands

Nitrogen (N) deposition continues to threaten upland ecosystems, contributing to acidification, eutrophication and biodiversity loss. We present results from a monitoring study aimed at investigating the fate of this deposited N within a pristine catchment in the Cairngorm Mountains (Scotland). Six sites were established along an elevation gradient (486-908 m) spanning the key habitats of temperate maritime uplands. Bulk deposition chemistry, soil carbon content, soil solution chemistry, soil temperature and soil moisture content were monitored over a 5 year period. Results were used to assess spatial variability in soil solution N and to investigate the factors and processes driving this variability. Highest soil solution inorganic N concentrations were found in the alpine soils at the top of the hillslope. Soil carbon stock, soil solution dissolved organic carbon (DOC) and factors representing site hydrology were the best predictors of NO(3)(-) concentration, with highest concentrations at low productivity sites with low DOC and freely-draining soils. These factors act as proxies for changing net biological uptake and soil/water contact time, and therefore support the hypothesis that spatial variations in soil solution NO(3)(-) are controlled by habitat N retention capacity. Soil percent carbon was a better predictor of soil solution inorganic N concentration than mass of soil carbon. NH(4)(+) was less affected by soil hydrology than NO(3)(-) and showed the effects of net mineralization inputs, particularly at Racomitrium heath and peaty sites. Soil solution dissolved organic N concentration was strongly related to both DOC and temperature, with a stronger temperature effect at more productive sites. Due to the spatial heterogeneity in N leaching potential, a fine-scale approach to assessing surface water vulnerability to N leaching is recommended over the broad scale, critical loads approach currently in use, particularly for sensitive areas.

Asynchrony among local communities stabilises ecosystem function of metacommunities

Abstract Temporal stability of ecosystem functioning increases the predictability and reliability of ecosystem services, and understanding the drivers of stability across spatial scales is important for land management and policy decisions. We used species‐level abundance data from 62 plant communities across five continents to assess mechanisms of temporal stability across spatial scales. We assessed how asynchrony (i.e. different units responding dissimilarly through time) of species and local communities stabilised metacommunity ecosystem function. Asynchrony of species increased stability of local communities, and asynchrony among local communities enhanced metacommunity stability by a wide range of magnitudes (1–315%); this range was positively correlated with the size of the metacommunity. Additionally, asynchronous responses among local communities were linked with species’ populations fluctuating asynchronously across space, perhaps stemming from physical and/or competitive differences among local communities. Accordingly, we suggest spatial heterogeneity should be a major focus for maintaining the stability of ecosystem services at larger spatial scales.

Developing monitoring protocols for cost-effective surveillance of lichens

The criteria set out by the International Union for Conservation of Nature to identify threatened species requires information on population trends which, for priority lichen species within Scotland, is lacking. Collecting such data is problematic as there is a lack of empirical information on the performance of different sampling designs and survey methodologies. Using Pseudocyphellaria norvegica as an example species, we tested differences in the efficiency of 3 transect patterns and a 20 minute search for surveying 100×100 m cells of potentially suitable habitat. The methods were not intended to census the total population of the cells but, rather, to provide a standardized, repeatable estimate of the population density to allow detection of trends through time. We also tested the repeatability of the methods between surveyors. The results provided no evidence to suggest that controlled survey methodologies using fixed transect patterns were any better in terms of consistency between surveyors or numbers of occupied trees found than 20 minute searches of the areas within each 100×100 m cell deemed suitable for the target species by an experienced surveyor. Given that following the fixed transect patterns took approximately twice as long as a 20 minute search, the search method would clearly be more cost-effective when there are large numbers of cells to survey. For all survey methods variability between surveyors was high, meaning that it would be extremely difficult to detect temporal changes in populations, and hence identify population trends. We also examined the extent to which recording presence/absence at the 1 ha scale might improve consistency between surveyors and found that it reduced, but did not eliminate, the surveyor variability. Recording only presence/absence would allow greater numbers of cells to be surveyed using the same level of resources, but would reduce the amount of information available per cell for use in analysis of population trends. We conclude that controlling inter-surveyor variability while collecting adequate data for population trend analysis is a major issue when planning and implementing any large-scale survey of lichen species.

Relative importance of local- and large-scale drivers of alpine soil microarthropod communities

Nitrogen (N) deposition and climate are acknowledged drivers of change in biodiversity and ecosystem function at large scales. However, at a local scale, their impact on functions and community structure of organisms is filtered by drivers like habitat quality and food quality/availability. This study assesses the relative impact of large-scale factors, N deposition and climate (rainfall and temperature), versus local-scale factors of habitat quality and food quality/availability on soil fauna communities at 15 alpine moss–sedge heaths along an N deposition gradient in the UK. Habitat quality and food quality/availability were the primary drivers of microarthropod communities. No direct impacts of N deposition on the microarthropod community were observed, but induced changes in habitat quality (decline in moss cover and depth) and food quality (decreased vegetation C:N) associated with increased N deposition strongly suggest an indirect impact of N. Habitat quality and climate explained variation in the composition of the Oribatida, Mesostigmata, and Collembola communities, while only habitat quality significantly impacted the Prostigmata. Food quality and prey availability were important in explaining the composition of the oribatid and mesostigmatid mite communities, respectively. This study shows that, in alpine habitats, soil microarthropod community structure responds most strongly to local-scale variation in habitat quality and food availability rather than large-scale variation in climate and pollution. However, given the strong links between N deposition and the key habitat quality parameters, we conclude that N deposition indirectly drives changes in the soil microarthropod community, suggesting a mechanism by which large-scale drivers indirectly impacts these functionally important groups.