[Show abstract][Hide abstract] ABSTRACT: Nutrient pollution presents a serious threat to biodiversity conservation. In terrestrial ecosystems, the deleterious effects of nitrogen pollution are increasingly understood and several mitigating environmental policies have been developed. Compared to nitrogen, the effects of increased phosphorus have received far less attention, although some studies have indicated that phosphorus pollution may be detrimental for biodiversity as well. Based on a dataset covering 501 grassland plots throughout Europe, we demonstrate that, independent of the level of atmospheric nitrogen deposition and soil acidity, plant species richness was consistently negatively related to soil phosphorus. We also identified thresholds in soil phosphorus above which biodiversity appears to remain at a constant low level. Our results indicate that nutrient management policies biased towards reducing nitrogen pollution will fail to preserve biodiversity. As soil phosphorus is known to be extremely persistent and we found no evidence for a critical threshold below which no environmental harm is expected, we suggest that agro-environmental schemes should include grasslands that are permanently free from phosphorus fertilization. This article is protected by copyright. All rights reserved.
Global Change Biology 06/2014; · 8.22 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: This chapter reports the findings of a Working Group on how atmospheric nitrogen (N) deposition affects both terrestrial and freshwater biodiversity. Regional and global scale impacts on biodiversity are addressed, together with potential indicators. Key conclusions are that: the rates of loss in biodiversity are greatest at the lowest and initial stages of N deposition increase; changes in species compositions are related to the relative amounts of N, carbon (C) and phosphorus (P) in the plant soil system; enhanced N inputs have implications for C cycling; N deposition is known to be having adverse effects on European and North American vegetation composition; very little is known about tropical ecosystem responses, while tropical ecosystems are major biodiversity hotspots and are increasingly recipients of very high N deposition rates; N deposition alters forest fungi and mycorrhyzal relations with plants; the rapid response of forest fungi and arthropods makes them good indicators of change; predictive tools (models) that address ecosystem scale processes are necessary to address complex drivers and responses, including the integration of N deposition, climate change and land use effects; criteria can be identified for projecting sensitivity of terrestrial and aquatic ecosystems to N deposition. Future research and policy-relevant recommendations are identified.
[Show abstract][Hide abstract] ABSTRACT: Humans are both intentionally (fertilization) and unintentionally (atmospheric nutrient deposition) adding nutrients worldwide. Increasing availability of biologically reactive nitrogen (N) is one of the major drivers of plant species loss. It remains unclear, however, whether plant diversity will be equally reduced by inputs of reactive N coming from either small and frequent N deposition events or large and infrequent N fertilization events. By independently manipulating the rate and frequency of reactive N inputs, our study teases apart these potentially contrasting effects. Plant species richness decreased more quickly at high rates and at low frequency of N addition, which suggests that previous fertilization studies have likely over-estimated the effects of N deposition on plant species loss. N-induced species loss resulted from both acidification and ammonium toxicity. Further study of small and frequent N additions will be necessary to project future rates of plant species loss under increasing aerial N deposition.This article is protected by copyright. All rights reserved.
Global Change Biology 04/2014; · 8.22 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Studies of experimental grassland communities have demonstrated that plant diversity can stabilize productivity through species asynchrony, in which decreases in the biomass of some species are compensated for by increases in others. However, it remains unknown whether these findings are relevant to natural ecosystems, especially those for which species diversity is threatened by anthropogenic global change. Here we analyse diversity-stability relationships from 41 grasslands on five continents and examine how these relationships are affected by chronic fertilization, one of the strongest drivers of species loss globally. Unmanipulated communities with more species had greater species asynchrony, resulting in more stable biomass production, generalizing a result from biodiversity experiments to real-world grasslands. However, fertilization weakened the positive effect of diversity on stability. Contrary to expectations, this was not due to species loss after eutrophication but rather to an increase in the temporal variation of productivity in combination with a decrease in species asynchrony in diverse communities. Our results demonstrate separate and synergistic effects of diversity and eutrophication on stability, emphasizing the need to understand how drivers of global change interactively affect the reliable provisioning of ecosystem services in real-world systems.
[Show abstract][Hide abstract] ABSTRACT: The global nitrogen (N) cycle has been transformed by human use of reactive N as a consequence of increased demand for food and energy. Given the considerable impact of humans on the N cycle, it is essential that we raise awareness amongst the public and policy makers as this is the first step in providing individuals and governments the opportunity to reduce their impact on the N cycle and reduce the environmental and health consequences of N pollution. Here we describe an N footprint tool for the UK developed as part of the N-PRINT program. The current per capita N footprint in the UK is 27.1 kg N per capita per year with food production constituting the largest proportion of the footprint (18.0 kg N per capita per year). Calculating an N footprint for 1971 (26.0 kg N per capita per year) demonstrates that per capita N footprints have increased slightly. The average UK footprint is smaller than that found in the USA but is higher than the Netherlands and Germany. Scenario analysis demonstrates that reducing food protein consumption to the levels recommended by the FAO and World Health Organization reduces the overall N footprint by 33%. Consuming a vegetarian diet and consuming only sustainable food both decreased the N footprint by 15% but changes in energy use have a much smaller impact.
[Show abstract][Hide abstract] ABSTRACT: Aim Evidence linking the accumulation of exotic species to the suppression of native diversity is equivocal, often relying on data from studies that have used different methods. Plot-level studies often attribute inverse relationships between native and exotic diversity to competition, but regional abiotic filters, including anthropogenic influences, can produce similar patterns. We seek to test these alternatives using identical scale-dependent sampling protocols in multiple grasslands on two continents. Location Thirty-two grassland sites in North America and Australia. Methods We use multiscale observational data, collected identically in grain and extent at each site, to test the association of local and regional factors with the plot-level richness and abundance of native and exotic plants. Sites captured environmental and anthropogenic gradients including land-use intensity, human population density, light and soil resources, climate and elevation. Site selection occurred independently of exotic diversity, meaning that the numbers of exotic species varied randomly thereby reducing potential biases if only highly invaded sites were chosen. Results Regional factors associated directly or indirectly with human activity had the strongest associations with plot-level diversity. These regional drivers had divergent effects: urban-based economic activity was associated with high exotic : native diversity ratios; climate- and landscape-based indicators of lower human population density were associated with low exotic : native ratios. Negative correlations between plot-level native and exotic diversity, a potential signature of competitive interactions, were not prevalent; this result did not change along gradients of productivity or heterogeneity. Main conclusion We show that plot-level diversity of native and exotic plants are more consistently associated with regional-scale factors relating to urbanization and climate suitability than measures indicative of competition. These findings clarify the long-standing difficulty in resolving drivers of exotic diversity using single-factor mechanisms, suggesting that multiple interacting anthropogenic-based processes best explain the accumulation of exotic diversity in modern landscapes.
[Show abstract][Hide abstract] ABSTRACT: Human alterations to nutrient cycles1, 2 and herbivore communities3, 4, 5, 6, 7 are affecting global biodiversity dramatically2. Ecological theory predicts these changes should be strongly counteractive: nutrient addition drives plant species loss through intensified competition for light, whereas herbivores prevent competitive exclusion by increasing ground-level light, particularly in productive systems8, 9. Here we use experimental data spanning a globally relevant range of conditions to test the hypothesis that herbaceous plant species losses caused by eutrophication may be offset by increased light availability due to herbivory. This experiment, replicated in 40 grasslands on 6 continents, demonstrates that nutrients and herbivores can serve as counteracting forces to control local plant diversity through light limitation, independent of site productivity, soil nitrogen, herbivore type and climate. Nutrient addition consistently reduced local diversity through light limitation, and herbivory rescued diversity at sites where it alleviated light limitation. Thus, species loss from anthropogenic eutrophication can be ameliorated in grasslands where herbivory increases ground-level light.
[Show abstract][Hide abstract] ABSTRACT: We aimed to answer the question of whether the species richness and composition of calcareous grasslands in North-western Germany had changed over the last 70 years as a result of atmospheric nitrogen (N) deposition. In total, 1186 plots of Festuco-Brometea (alliance Bromion erecti) grasslands from the sub-oceanic regions of the country were compiled (1061 plots from literature sources spanning a time period from 1936 to 1996, 125 new plots from 2008). Environmental descriptors recorded for each plot included geographic coordinates, altitude, heat index (combining slope and aspect), mean Ellenberg indicator values for light, soil moisture, soil pH and soil N, and cumulative N deposition (the latter being highly positively correlated with the year of sampling).
In a Detrended Correspondence Analysis, the sample plot scores along axis one were highly correlated with the mean Ellenberg N-values, those along axis two were significantly affected by the year of sampling. In a general linear model, species richness of vascular plants showed a markedly hump-shaped relationship with mean Ellenberg N-value, whereas it was weakly affected by year (cumulative N load). Species with a significant negative trend over time were more often (than expected by chance) habitat specialists of dry grasslands, small, light-demanding and winter-green or evergreen with smaller seeds and scleromorphic leaves. In contrast to what has been found for acidic grasslands, N deposition in calcareous grasslands did not result in a decline in species richness, most likely because calcareous grasslands are water- and phosphorus-limited, and are well-buffered in terms of soil pH. To prevent a further change in species composition towards more mesophytic communities, grassland management by the site managers needs to be intensified.
[Show abstract][Hide abstract] ABSTRACT: Invasions have increased the size of regional species pools, but are typically assumed to reduce native diversity. However, global-scale tests of this assumption have been elusive because of the focus on exotic species richness, rather than relative abundance. This is problematic because low invader richness can indicate invasion resistance by the native community or, alternatively, dominance by a single exotic species. Here, we used a globally-replicated study to quantify relationships between exotic richness and abundance in grass-dominated ecosystems in 13 countries on six continents, ranging from salt marshes to alpine tundra. We tested effects of human land use, native community diversity, herbivore pressure, and nutrient limitation on exotic plant dominance. Despite its widespread use, exotic richness was a poor proxy for exotic dominance at low exotic richness, because sites that contained few exotic species ranged from relatively pristine (low exotic richness and cover) to almost completely exotic-dominated (low exotic richness but high exotic cover). Both exotic cover and richness were predicted by native plant diversity (native grass richness) and land use (distance to cultivation). Although climate was important for predicting both exotic cover and richness, climatic factors predicting cover (precipitation variability) differed from those predicting richness (maximum temperature and temperature in the wettest quarter). Herbivory and nutrient limitation did not predict exotic richness or cover. Exotic dominance varied most among regions (subcontinents), whereas cover was greatest in areas with low native grass richness at the site- or regional-scale. Although this could reflect native grass displacement, a lack of biotic resistance is a more likely explanation, given that grasses comprise the most aggressive invaders. These findings underscore the need to move beyond richness as a surrogate for the extent of invasion, because this metric confounds mono-dominance with invasion resistance. Monitoring species' relative abundance will more rapidly advance our understanding of invasions. This article is protected by copyright. All rights reserved.
Global Change Biology 08/2013; · 8.22 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: N deposition is currently affecting nutrient cycling in terrestrial ecosystems. We studied the effects of four years of N application (0, 10, 20 and 50kgNha(-1)year(-1)+background deposition) on soil chemistry and fertility in a semiarid shrubland in central Spain. Soil pH and nutrient availability fluctuated seasonally. The inorganic-N fraction in soil was dominated by nitrate, as expected in calcareous soils. N application increased inorganic N availability in soil. There was a negative correlation between N application and soil K(+) availability and pH, measured as the % change after four years. Soil N and C storage (evaluated as the % change) slightly increased after four years. Our data suggest that, in the short-term, the seasonality of nutrients overwhelm any chemical alteration related to N deposition. However, the potential implication of continuous N addition on soil chemistry in the long-term is not well understood.
Science of The Total Environment 03/2013; 452-453C:78-86. · 3.26 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Chronically elevated reactive nitrogen deposition has a severe impact on many ecosystems, and there is widespread interest in the possibility of using plant community composition to estimate the level of nitrogen deposition and consequent impacts. Existing approaches use a variety of simple measures including functional type ratios, Ellenberg numbers, and diversity indices. We propose an alternative approach in which species–environment models are constructed using national datasets designed to capture broad-scale deposition patterns. We construct models using partial least squares, weighted average, and maximum likelihood Gaussian logit regression for two British semi-natural habitats, and test how well they predict N deposition by cross-validation. We find that performance is good with R2 values up to 0.7, and suggest that such models could be a useful addition to the bioindication toolbox.
[Show abstract][Hide abstract] ABSTRACT: Plant growth can be limited by resource acquisition and defence against consumers, leading to contrasting trade-off possibilities. The competition-defence hypothesis posits a trade-off between competitive ability and defence against enemies (e.g. herbivores and pathogens). The growth-defence hypothesis suggests that strong competitors for nutrients are also defended against enemies, at a cost to growth rate. We tested these hypotheses using observations of 706 plant populations of over 500 species before and following identical fertilisation and fencing treatments at 39 grassland sites worldwide. Strong positive covariance in species responses to both treatments provided support for a growth-defence trade-off: populations that increased with the removal of nutrient limitation (poor competitors) also increased following removal of consumers. This result held globally across 4 years within plant life-history groups and within the majority of individual sites. Thus, a growth-defence trade-off appears to be the norm, and mechanisms maintaining grassland biodiversity may operate within this constraint.
[Show abstract][Hide abstract] ABSTRACT: Based on regional-scale studies, aboveground production and litter decomposition are thought to positively covary, because they are driven by shared biotic and climatic factors. Until now we have been unable to test whether production and decomposition are generally coupled across climatically dissimilar regions, because we lacked replicated data collected within a single vegetation type across multiple regions, obfuscating the drivers and generality of the association between production and decomposition. Furthermore, our understanding of the relationships between production and decomposition rests heavily on separate meta-analyses of each response, because no studies have simultaneously measured production and the accumulation or decomposition of litter using consistent methods at globally relevant scales. Here, we use a multi-country grassland dataset collected using a standardized protocol to show that live plant biomass (an estimate of aboveground net primary production) and litter disappearance (represented by mass loss of aboveground litter) do not strongly covary. Live biomass and litter disappearance varied at different spatial scales. There was substantial variation in live biomass among continents, sites and plots whereas among continent differences accounted for most of the variation in litter disappearance rates. Although there were strong associations among aboveground biomass, litter disappearance and climatic factors in some regions (e.g. U.S. Great Plains), these relationships were inconsistent within and among the regions represented by this study. These results highlight the importance of replication among regions and continents when characterizing the correlations between ecosystem processes and interpreting their global-scale implications for carbon flux. We must exercise caution in parameterizing litter decomposition and aboveground production in future regional and global carbon models as their relationship is complex.
PLoS ONE 01/2013; 8(2):e54988. · 3.53 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: In Europe and, increasingly, the rest of the world, the key policy tool for the control of air pollution is the critical load, a level of pollution below which there are no known significant harmful effects on the environment. Critical loads are used to map sensitive regions and habitats, permit individual polluting activities, and frame international negotiations on transboundary air pollution. Despite their fundamental importance in environmental science and policy, there has been no systematic attempt to verify a critical load with field survey data. Here, we use a large dataset of European grasslands along a gradient of nitrogen (N) deposition to show statistically significant declines in the abundance of species from the lowest level of N deposition at which it is possible to identify a change. Approximately 60% of species change points occur at or below the range of the currently established critical load. If this result is found more widely, the underlying principle of no harm in pollution policy may need to be modified to one of informed decisions on how much harm is acceptable. Our results highlight the importance of protecting currently unpolluted areas from new pollution sources, because we cannot rule out ecological impacts from even relatively small increases in reactive N deposition.
Proceedings of the National Academy of Sciences 01/2013; 110(3):984-987. · 9.81 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Anthropogenically increased input of nitrogen (N) and phosphorous (P) have led to a severe reduction of plant species richness in European semi-natural grasslands. Although it is well established that this species loss is not trait neutral, a thorough analysis of the effects of nutrient addition on trait based func-tional diversity and functional composition, inde-pendently of species loss, is lacking so far. We compiled data on the plant species abundance (re-levé 's) of 279 Nardus grasslands from nine European countries, across a gradient of soil N and P content. Functional diversity (Petchy and Gaston's FDc, weighted FDc and quadratic entropy) and mean trait composition were calculated for each relevé , based on 21 functional traits. Differences in functional diversity and functional composition were related to differences in soil N, atmospheric N deposition, soil P and pH, while controlling for geographic location and species richness. All functional diversity mea-sures decreased with increasing soil N, with wFDc also decreased by soil P, independent of species loss. This was accompanied by clear shifts in functional trait composition, associated with shifts from below-ground competition for nutrients to above-ground competition for light. This resulted in a decrease in insect-pollinated therophytes and chamaephytes and an increase in long-lived, clonal graminoids and hemicryptophytes under increasing soil N and P. These functional community changes can be ex-pected to alter both ecosystem functioning and ser-vice provisioning of the studied grasslands. Our research emphasizes the importance of a reduction of both N and P emission throughout Europe for sustainable conservation of these communities.
[Show abstract][Hide abstract] ABSTRACT: Emissions and deposition of ammonia and nitrogen oxides have strongly increased since the 1950s. This has led to significant changes in the nitrogen (N) cycle, vegetation composition and plant diversity in many ecosystems of high conservation value in Europe. As a consequence of different regional pollution levels and of the increased importance of reduced N in the near future, determining the effect of different forms of N is an important task for understanding the consequences of atmospheric N inputs. We have initiated three replicated N addition experiments in species-rich, acidic grasslands spanning a climatic gradient in the Atlantic biogeographic region of Europe in Norway, Wales and France at sites with low levels of pollution. N was added in two doses (0 and 70 kg N ha−1 year−1 above background) and in three forms (oxidised N, reduced N and a 50–50 combination). After 2.5 years of N additions, the effects of these treatments on plant biomass, plant nutritional status, soil pH and soil nutrient availability were determined. Impacts of the N additions were observed within the 2.5-year research period. In some cases, the first signs of differential effects of N form could also be demonstrated. In the French site, for example, grass biomass was significantly increased by the oxidised N treatments but decreased by the reduced N treatments. In the Norwegian site, the reduced N treatments significantly reduced soil pH, whereas oxidised N did not. Effects on nutrient availability were also observed. These experiments will be continued to elucidate the longer term impacts of N deposition on these grasslands.
Water Air and Soil Pollution 01/2013; 224(9). · 1.75 Impact Factor