Global change effects on plant communities are magnified by time and the number of global change factors imposed.

Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (<10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity-ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously.

Determinants of community compositional change are equally affected by global change.

Global change is impacting plant community composition, but the mechanisms underlying these changes are unclear. Using a dataset of 58 global change experiments, we tested the five fundamental mechanisms of community change: changes in evenness and richness, reordering, species gains and losses. We found 71% of communities were impacted by global change treatments, and 88% of communities that were exposed to two or more global change drivers were impacted. Further, all mechanisms of change were equally likely to be affected by global change treatments-species losses and changes in richness were just as common as species gains and reordering. We also found no evidence of a progression of community changes, for example, reordering and changes in evenness did not precede species gains and losses. We demonstrate that all processes underlying plant community composition changes are equally affected by treatments and often occur simultaneously, necessitating a wholistic approach to quantifying community changes.

Legacy effects of nitrogen and phosphorus additions on vegetation and carbon stocks of upland heaths.

Soil carbon (C) pools and plant community composition are regulated by nitrogen (N) and phosphorus (P) availability. Atmospheric N deposition impacts ecosystem C storage, but the direction of response varies between systems. Phosphorus limitation may constrain C storage response to N, hence P application to increase plant productivity and thus C sequestration has been suggested. We revisited a 23-yr-old field experiment where N and P had been applied to upland heath, a widespread habitat supporting large soil C stocks. At 10 yr after the last nutrient application we quantified long-term changes in vegetation composition and in soil and vegetation C and P stocks. Nitrogen addition, particularly when combined with P, strongly influenced vegetation composition, favouring grasses over Calluna vulgaris, and led to a reduction in vegetation C stocks. However, soil C stocks did not respond to nutrient treatments. We found 40% of the added P had accumulated in the soil. This study showed persistent effects of N and N + P on vegetation composition, whereas effects of P alone were small and showed recovery. We found no indication that P application could mitigate the effects of N on vegetation or increase C sequestration in this system.

Nitrogen deposition drives loss of moss cover in alpine moss-sedge heath via lowered C : N ratio and accelerated decomposition.

In alpine ecosystems, nitrogen (N) deposition has been linked to plant community composition change, including loss of bryophytes and increase of graminoids. Since bryophyte growth is stimulated by increased N availability, it has been hypothesized that loss of bryophyte cover is driven by enhanced decomposition. As bryophyte mats are a significant carbon (C) store, their loss may impact C storage in these ecosystems. We used an N deposition gradient across 15 sites in the UK to examine effects of N deposition on bryophyte litter quality, decomposition and C and N stocks in Racomitrium moss-sedge heath. Increasing N deposition reduced C : N in bryophyte litter, which in turn enhanced decomposition. Soil N stocks increased significantly in response to increased N deposition, and soil C : N declined. However, depletion of the bryophyte mat and its replacement by graminoids under high N deposition was not associated with a change in total ecosystem C stocks. We conclude that decomposition processes in Racomitrium heath are very sensitive to N deposition and provide a mechanism by which N deposition drives depletion of the bryophyte mat. Nitrogen deposition did not measurably alter C stocks, but changes in soil N stocks and C : N suggest the ecosystem is becoming N saturated.

Asynchrony among local communities stabilises ecosystem function of metacommunities.

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.

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.

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.

Impacts of nitrogen deposition on carbon and nitrogen cycling in alpine Racomitrium heath in the UK and prospects for recovery.

Deposition of reactive nitrogen (N) is a major threat to terrestrial ecosystems associated with impacts on ecosystem properties and functions including carbon (C) and nutrient stocks, soil water quality and nutrient retention. In the oceanic-alpine Racomitrium heath habitat, N deposition is associated with moss mat degradation and a shift from bryophyte to graminoid dominance. To investigate the effects of moss mat decline on C and N stocks and fluxes, we collected Racomitrium heath vegetation/soil cores from sites along a gradient of N deposition in the UK. Cores were maintained under controlled conditions and exposed to scenarios of current (8-40 kg N ha-1 y-1), reduced (8 kg N ha-1 y-1) and elevated (50 kg N ha-1 y-1) N deposition. Cores from high N deposition sites had smaller aboveground C and N stocks and, under current conditions, leached large amounts of inorganic N and had low soil water pH compared with low N deposition sites. With reduced N deposition there was evidence for rapid recovery of soil water quality in terms of reduced N leaching and small increases in pH. Under high N deposition, cores from low N deposition sites retained much of the applied N while those with a history of high N deposition leached large amounts of inorganic N. Carbon fluxes in soil water and net CO2 fluxes varied according to core source site but were not affected by the N deposition scenarios. We conclude that C and N stocks and cycling in Racomitrium heath are strongly affected by long-term exposure to N deposition but that soil water quality may improve rapidly, if N deposition rates are reduced. The legacy of N deposition impacts on moss mat cover and vegetation composition however, mean that the ecosystem remains sensitive to future pulses in N input.

The potential for modelling peatland habitat condition in Scotland using long-term MODIS data.

Globally, peatlands provide an important sink of carbon in their near natural state but potentially act as a source of gaseous and dissolved carbon emission if not in good condition. There is a pressing need to remotely identify peatland sites requiring improvement and to monitor progress following restoration. A medium resolution model was developed based on a training dataset of peatland habitat condition and environmental covariates, such as morphological features, against information derived from the Moderate Resolution Imaging Spectroradiometer (MODIS), covering Scotland (UK). The initial, unrestricted, model provided the probability of a site being in favourable condition. Receiver operator characteristics (ROC) curves for restricted training data, limited to those located on a peat soil map, resulted in an accuracy of 0.915. The kappa statistic was 0.8151, suggesting good model fit. The derived map of predicted peatland condition at the suggested 0.56 threshold was corroborated by data from other sources, including known restoration sites, areas under known non-peatland land cover and previous vegetation survey data mapped onto inferred condition categories. The resulting locations of the areas of peatland modelled to be in favourable ecological condition were largely confined to the North and West of the country, which not only coincides with prior land use intensity but with published predictions of future retraction of the bioclimatic space for peatlands. The model is limited by a lack of spatially appropriate ground observations, and a lack of verification of peat depth at training site locations, hence future efforts to remotely assess peatland condition will require more appropriate ground-based monitoring. If appropriate ground-based observations could be collected, using remote sensing could be considered a cost-efficient means to provide data on changes in peatland habitat condition.

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.

Growth responses of low-alpine dwarf-shrub heath species to nitrogen deposition and management.

Nitrogen deposition is a continuing problem in European alpine regions. We hypothesised that, despite climatic limitations, low-alpine Calluna heathland would respond to nitrogen addition with increased shoot growth and flowering and that fire and grazing would modify responses. In a five-year study, 0-50kgNha(-1)y(-1) were added, combined with burning (+/-) and clipping (+/-). Calluna vulgaris responded with increased shoot extension, but effects on flowering were variable. Burning enhanced the positive effect of nitrogen addition and negative effects of clipping. Sub-dominant shrubs generally did not respond to nitrogen. C. vulgaris shoot extension was stimulated by nitrogen addition of 10kgNha(-1)y(-1) (above background) supporting suggestions that alpine heathlands are sensitive to low levels of nitrogen deposition. Increased C. vulgaris growth could negatively impact on important lichen components of this vegetation through increased shading and competition. Climatic factors constrain productivity in this community, but do not prevent rapid responses to nitrogen deposition by some species.

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.

Assessing the recovery potential of alpine moss-sedge heath: reciprocal transplants along a nitrogen deposition gradient.

The potential of alpine moss-sedge heath to recover from elevated nitrogen (N) deposition was assessed by transplanting Racomitrium lanuginosum shoots and vegetation turfs between 10 elevated N deposition sites (8.2-32.9 kg ha(-1) yr(-1)) and a low N deposition site, Ben Wyvis (7.2 kg ha(-1) yr(-1)). After two years, tissue N of Racomitrium shoots transplanted from higher N sites to Ben Wyvis only partially equilibrated to reduced N deposition whereas reciprocal transplants almost matched the tissue N of indigenous moss. Unexpectedly, moss shoot growth was stimulated at higher N deposition sites. However, moss depth and biomass increased in turfs transplanted to Ben Wyvis, apparently due to slower shoot turnover (suggested to result partly from decreased tissue C:N slowing decomposition), whilst abundance of vascular species declined. Racomitrium heath has the potential to recover from the impacts of N deposition; however, this is constrained by the persistence of enhanced moss tissue N contents.

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.