Publications by authors named "Carly J Stevens"

45 Publications

Comment on Pescott & Jitlal 2020: Failure to account for measurement error undermines their conclusion of a weak impact of nitrogen deposition on plant species richness.

PeerJ 2021 12;9:e10632. Epub 2021 Jan 12.

Land Use, Centre for Ecology & Hydrology Lancaster, Lancaster, United Kingdom.

Estimation of the impacts of atmospheric nitrogen (N) deposition on ecosystems and biodiversity is a research imperative. Analyses of large-scale spatial gradients, where an observed response is correlated with measured or modelled deposition, have been an important source of evidence. A number of problems beset this approach. For example, if responses are spatially aggregated then treating each location as statistically independent can lead to biased confidence intervals and a greater probably of false positive results. Using methods that account for residual spatial autocorrelation, Pescott & Jitlal (2020) re-analysed two large-scale spatial gradient datasets from Britain where modelled N deposition at 5 × 5 km resolution had been previously correlated with species richness in small quadrats. They found that N deposition effects were weaker than previously demonstrated leading them to conclude that ". We use a simulation study to show that their conclusion is unreliable despite them recognising that an influential fraction of the residual spatially structured variation could itself be attributable to N deposition. This arises because the covariate used was modelled N deposition at 5 × 5 km resolution leaving open the possibility that measured or modelled N deposition at finer resolutions could explain more variance in the response. Explicitly treating this as spatially auto-correlated error ignores this possibility and leads directly to their unreliable conclusion. We further demonstrate the plausibility of this scenario by showing that significant variation in N deposition at the 1 km square resolution is indeed averaged at 5 × 5 km resolution. Further analyses are required to explore whether estimation of the size of the N deposition effect on plant species richness and other measures of biodiversity is indeed dependent on the accuracy and hence measurement error of the N deposition covariate. Until then the conclusions of Pescott & Jitlal (2020) should be considered premature.
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http://dx.doi.org/10.7717/peerj.10632DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7810039PMC
January 2021

Increasing effects of chronic nutrient enrichment on plant diversity loss and ecosystem productivity over time.

Ecology 2021 02 18;102(2):e03218. Epub 2021 Jan 18.

Department of Ecology, Evolution, and Behavior, University of Minnesota, St. Paul, Minnesota, 55108, USA.

Human activities are enriching many of Earth's ecosystems with biologically limiting mineral nutrients such as nitrogen (N) and phosphorus (P). In grasslands, this enrichment generally reduces plant diversity and increases productivity. The widely demonstrated positive effect of diversity on productivity suggests a potential negative feedback, whereby nutrient-induced declines in diversity reduce the initial gains in productivity arising from nutrient enrichment. In addition, plant productivity and diversity can be inhibited by accumulations of dead biomass, which may be altered by nutrient enrichment. Over longer time frames, nutrient addition may increase soil fertility by increasing soil organic matter and nutrient pools. We examined the effects of 5-11 yr of nutrient addition at 47 grasslands in 12 countries. Nutrient enrichment increased aboveground live biomass and reduced plant diversity at nearly all sites, and these effects became stronger over time. We did not find evidence that nutrient-induced losses of diversity reduced the positive effects of nutrients on biomass; however, nutrient effects on live biomass increased more slowly at sites where litter was also increasing, regardless of plant diversity. This work suggests that short-term experiments may underestimate the long-term nutrient enrichment effects on global grassland ecosystems.
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http://dx.doi.org/10.1002/ecy.3218DOI Listing
February 2021

Global impacts of fertilization and herbivore removal on soil net nitrogen mineralization are modulated by local climate and soil properties.

Glob Chang Biol 2020 Dec 22;26(12):7173-7185. Epub 2020 Sep 22.

Grupo de Investigaciones en Biología de la Conservación, INIBIOMA (CONICET-UNCOMA), Bariloche, Argentina.

Soil nitrogen (N) availability is critical for grassland functioning. However, human activities have increased the supply of biologically limiting nutrients, and changed the density and identity of mammalian herbivores. These anthropogenic changes may alter net soil N mineralization (soil net N ), that is, the net balance between N mineralization and immobilization, which could severely impact grassland structure and functioning. Yet, to date, little is known about how fertilization and herbivore removal individually, or jointly, affect soil net N across a wide range of grasslands that vary in soil and climatic properties. Here we collected data from 22 grasslands on five continents, all part of a globally replicated experiment, to assess how fertilization and herbivore removal affected potential (laboratory-based) and realized (field-based) soil net N . Herbivore removal in the absence of fertilization did not alter potential and realized soil net N . However, fertilization alone and in combination with herbivore removal consistently increased potential soil net N Realized soil net N , in contrast, significantly decreased in fertilized plots where herbivores were removed. Treatment effects on potential and realized soil net N were contingent on site-specific soil and climatic properties. Fertilization effects on potential soil net N were larger at sites with higher mean annual precipitation (MAP) and temperature of the wettest quarter (T.q.wet). Reciprocally, realized soil net N declined most strongly with fertilization and herbivore removal at sites with lower MAP and higher T.q.wet. In summary, our findings show that anthropogenic nutrient enrichment, herbivore exclusion and alterations in future climatic conditions can negatively impact soil net N across global grasslands under realistic field conditions. This is an important context-dependent knowledge for grassland management worldwide.
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http://dx.doi.org/10.1111/gcb.15308DOI Listing
December 2020

Nutrient availability controls the impact of mammalian herbivores on soil carbon and nitrogen pools in grasslands.

Glob Chang Biol 2020 Feb 3. Epub 2020 Feb 3.

Department of Terrestrial Ecology, Netherlands Institute of Ecology (NIOO-KNAW), Wageningen, The Netherlands.

Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature - herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local-scale herbivory, and its interaction with nutrient enrichment and climate, within global-scale models to better predict land-atmosphere interactions under future climate change.
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http://dx.doi.org/10.1111/gcb.15023DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7155038PMC
February 2020

Potential vulnerability of 348 herbaceous species to atmospheric deposition of nitrogen and sulfur in the United States.

Nat Plants 2019 07 1;5(7):697-705. Epub 2019 Jul 1.

Department of Botany, University of Wisconsin, Madison, WI, USA.

Atmospheric nitrogen and sulfur pollution increased over much of the United States during the twentieth century from fossil fuel combustion and industrial agriculture. Despite recent declines, nitrogen and sulfur deposition continue to affect many plant communities in the United States, although which species are at risk remains uncertain. We used species composition data from >14,000 survey sites across the contiguous United States to evaluate the association between nitrogen and sulfur deposition and the probability of occurrence for 348 herbaceous species. We found that the probability of occurrence for 70% of species was negatively associated with nitrogen or sulfur deposition somewhere in the contiguous United States (56% for N, 51% for S). Of the species, 15% and 51% potentially decreased at all nitrogen and sulfur deposition rates, respectively, suggesting thresholds below the minimum deposition they receive. Although more species potentially increased than decreased with nitrogen deposition, increasers tended to be introduced and decreasers tended to be higher-value native species. More vulnerable species tended to be shorter with lower tissue nitrogen and magnesium. These relationships constitute predictive equations to estimate critical loads. These results demonstrate that many herbaceous species may be at risk from atmospheric deposition and can inform improvements to air quality policies in the United States and globally.
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http://dx.doi.org/10.1038/s41477-019-0442-8DOI Listing
July 2019

Correction to: Mowing mitigates the negative impacts of N addition on plant species diversity.

Oecologia 2019 May;190(1):269-272

Erguna Forest-Steppe Ecotone Research Station, CAS Key Laboratory of Forest Ecology and Management, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang, 110016, China.

Unfortunately, the panels of (f) in Figures 1, 2, and 4.
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http://dx.doi.org/10.1007/s00442-019-04396-yDOI Listing
May 2019

Nitrogen in the environment.

Authors:
Carly J Stevens

Science 2019 Feb;363(6427):578-580

Lancaster Environment Centre, Lancaster University, Lancaster LA1 4YQ, UK.

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http://dx.doi.org/10.1126/science.aav8215DOI Listing
February 2019

Mowing mitigates the negative impacts of N addition on plant species diversity.

Oecologia 2019 Mar 6;189(3):769-779. Epub 2019 Feb 6.

Erguna Forest-Steppe Ecotone Research Station, CAS Key Laboratory of Forest Ecology and Management, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang, 110016, China.

Increasing availability of reactive nitrogen (N) threatens plant diversity in diverse ecosystems. While there is mounting evidence for the negative impacts of N deposition on one component of diversity, species richness, we know little about its effects on another one, species evenness. It is suspected that ecosystem management practice that removes nitrogen from the ecosystem, such as hay-harvesting by mowing in grasslands, would mitigate the negative impacts of N deposition on plant diversity. However, empirical evidence is scarce. Here, we reported the main and interactive effects of N deposition and mowing on plant diversity in a temperate meadow steppe with 4-year data from a field experiment within which multi-level N addition rates and multiple N compounds are considered. Across all the types of N compounds, species richness and evenness significantly decreased with the increases of N addition rate, which was mainly caused by the growth of a tall rhizomatous grass, Leymus chinensis. Such negative impacts of N addition were accumulating with time. Mowing significantly reduced the dominance of L. chinensis, and mitigated the negative impacts of N deposition on species evenness. We present robust evidence that N deposition threatened biodiversity by reducing both species richness and evenness, a process which could be alleviated by mowing. Our results highlight the changes of species evenness in driving the negative impacts of N deposition on plant diversity and the role of mowing in mediating such negative impacts of N deposition.
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http://dx.doi.org/10.1007/s00442-019-04353-9DOI Listing
March 2019

Leaf nutrients, not specific leaf area, are consistent indicators of elevated nutrient inputs.

Nat Ecol Evol 2019 03 4;3(3):400-406. Epub 2019 Feb 4.

Forest Research Centre, School of Agriculture, University of Lisbon, Lisbon, Portugal.

Leaf traits are frequently measured in ecology to provide a 'common currency' for predicting how anthropogenic pressures impact ecosystem function. Here, we test whether leaf traits consistently respond to experimental treatments across 27 globally distributed grassland sites across 4 continents. We find that specific leaf area (leaf area per unit mass)-a commonly measured morphological trait inferring shifts between plant growth strategies-did not respond to up to four years of soil nutrient additions. Leaf nitrogen, phosphorus and potassium concentrations increased in response to the addition of each respective soil nutrient. We found few significant changes in leaf traits when vertebrate herbivores were excluded in the short-term. Leaf nitrogen and potassium concentrations were positively correlated with species turnover, suggesting that interspecific trait variation was a significant predictor of leaf nitrogen and potassium, but not of leaf phosphorus concentration. Climatic conditions and pretreatment soil nutrient levels also accounted for significant amounts of variation in the leaf traits measured. Overall, we find that leaf morphological traits, such as specific leaf area, are not appropriate indicators of plant response to anthropogenic perturbations in grasslands.
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http://dx.doi.org/10.1038/s41559-018-0790-1DOI Listing
March 2019

What is the most ecologically-meaningful metric of nitrogen deposition?

Environ Pollut 2019 Apr 18;247:319-331. Epub 2019 Jan 18.

Centre for Ecology and Hydrology, Bush Estate, Penicuik, Midlothian, EH260QB, UK.

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.
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http://dx.doi.org/10.1016/j.envpol.2019.01.059DOI Listing
April 2019

Recent advances in understanding grasslands.

Authors:
Carly J Stevens

F1000Res 2018 30;7. Epub 2018 Aug 30.

Lancaster Environment Centre, Lancaster University, Lancaster, UK.

Grasslands are a vitally important ecosystem, supporting a wide range of ecosystem services and high levels of biodiversity. As a consequence, they have long been a focus for ecologists, playing host to some of the world's longest-running ecological experiments and providing the inspiration for many long-standing theories and debates. Because the field of grassland ecology is broad, encompassing many areas of ecology, this article picks some areas of particular debate and development to look at recent advances. The areas include relationships between diversity and productivity, ecosystem stability and ecosystem service provision, global change threats from nutrient addition, invasive species, climate change, and plant soil interactions.
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http://dx.doi.org/10.12688/f1000research.15050.1DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6117858PMC
July 2019

Spatial heterogeneity in species composition constrains plant community responses to herbivory and fertilisation.

Ecol Lett 2018 09 27;21(9):1364-1371. Epub 2018 Jun 27.

Department of Biology, Duke University, Durham, NC, 27708, USA.

Environmental change can result in substantial shifts in community composition. The associated immigration and extinction events are likely constrained by the spatial distribution of species. Still, studies on environmental change typically quantify biotic responses at single spatial (time series within a single plot) or temporal (spatial beta diversity at single time points) scales, ignoring their potential interdependence. Here, we use data from a global network of grassland experiments to determine how turnover responses to two major forms of environmental change - fertilisation and herbivore loss - are affected by species pool size and spatial compositional heterogeneity. Fertilisation led to higher rates of local extinction, whereas turnover in herbivore exclusion plots was driven by species replacement. Overall, sites with more spatially heterogeneous composition showed significantly higher rates of annual turnover, independent of species pool size and treatment. Taking into account spatial biodiversity aspects will therefore improve our understanding of consequences of global and anthropogenic change on community dynamics.
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http://dx.doi.org/10.1111/ele.13102DOI Listing
September 2018

Herbivory and eutrophication mediate grassland plant nutrient responses across a global climatic gradient.

Ecology 2018 04 31;99(4):822-831. Epub 2018 Mar 31.

Department of Ecology, Evolution, and Behavior, University of MN, St. Paul, Minnesota, 55108, USA.

Plant stoichiometry, the relative concentration of elements, is a key regulator of ecosystem functioning and is also being altered by human activities. In this paper we sought to understand the global drivers of plant stoichiometry and compare the relative contribution of climatic vs. anthropogenic effects. We addressed this goal by measuring plant elemental (C, N, P and K) responses to eutrophication and vertebrate herbivore exclusion at eighteen sites on six continents. Across sites, climate and atmospheric N deposition emerged as strong predictors of plot-level tissue nutrients, mediated by biomass and plant chemistry. Within sites, fertilization increased total plant nutrient pools, but results were contingent on soil fertility and the proportion of grass biomass relative to other functional types. Total plant nutrient pools diverged strongly in response to herbivore exclusion when fertilized; responses were largest in ungrazed plots at low rainfall, whereas herbivore grazing dampened the plant community nutrient responses to fertilization. Our study highlights (1) the importance of climate in determining plant nutrient concentrations mediated through effects on plant biomass, (2) that eutrophication affects grassland nutrient pools via both soil and atmospheric pathways and (3) that interactions among soils, herbivores and eutrophication drive plant nutrient responses at small scales, especially at water-limited sites.
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http://dx.doi.org/10.1002/ecy.2175DOI Listing
April 2018

Biogeographic differences in soil biota promote invasive grass response to nutrient addition relative to co-occurring species despite lack of belowground enemy release.

Oecologia 2018 03 5;186(3):611-620. Epub 2018 Feb 5.

Landcare Research, PO Box 69040, Lincoln, 7640, New Zealand.

Multiple plant species invasions and increases in nutrient availability are pervasive drivers of global environmental change that often co-occur. Many plant invasion studies, however, focus on single-species or single-mechanism invasions, risking an oversimplification of a multifaceted process. Here, we test how biogeographic differences in soil biota, such as belowground enemy release, interact with increases in nutrient availability to influence invasive plant growth. We conducted a greenhouse experiment using three co-occurring invasive grasses and one native grass. We grew species in live and sterilized soil from the invader's native (United Kingdom) and introduced (New Zealand) ranges with a nutrient addition treatment. We found no evidence for belowground enemy release. However, species' responses to nutrients varied, and this depended on soil origin and sterilization. In live soil from the introduced range, the invasive species Lolium perenne L. responded more positively to nutrient addition than co-occurring invasive and native species. In contrast, in live soil from the native range and in sterilized soils, there were no differences in species' responses to nutrients. This suggests that the presence of soil biota from the introduced range allowed L. perenne to capture additional nutrients better than co-occurring species. Considering the globally widespread nature of anthropogenic nutrient additions to ecosystems, this effect could be contributing to a global homogenization of flora and the associated losses in native species diversity.
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http://dx.doi.org/10.1007/s00442-018-4081-yDOI Listing
March 2018

Belowground competition drives invasive plant impact on native species regardless of nitrogen availability.

Oecologia 2018 02 7;186(2):577-587. Epub 2017 Dec 7.

Landcare Research, PO Box 69040, Lincoln, 7640, New Zealand.

Plant invasions and eutrophication are pervasive drivers of global change that cause biodiversity loss. Yet, how invasive plant impacts on native species, and the mechanisms underpinning these impacts, vary in relation to increasing nitrogen (N) availability remains unclear. Competition is often invoked as a likely mechanism, but the relative importance of the above and belowground components of this is poorly understood, particularly under differing levels of N availability. To help resolve these issues, we quantified the impact of a globally invasive grass species, Agrostis capillaris, on two co-occurring native New Zealand grasses, and vice versa. We explicitly separated above- and belowground interactions amongst these species experimentally and incorporated an N addition treatment. We found that competition with the invader had large negative impacts on native species growth (biomass decreased by half), resource capture (total N content decreased by up to 75%) and even nutrient stoichiometry (native species tissue C:N ratios increased). Surprisingly, these impacts were driven directly and indirectly by belowground competition, regardless of N availability. Higher root biomass likely enhanced the invasive grass's competitive superiority belowground, indicating that root traits may be useful tools for understanding invasive plant impacts. Our study shows that belowground competition can be more important in driving invasive plant impacts than aboveground competition in both low and high fertility ecosystems, including those experiencing N enrichment due to global change. This can help to improve predictions of how two key drivers of global change, plant species invasions and eutrophication, impact native species diversity.
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http://dx.doi.org/10.1007/s00442-017-4039-5DOI Listing
February 2018

Local loss and spatial homogenization of plant diversity reduce ecosystem multifunctionality.

Nat Ecol Evol 2018 Jan 4;2(1):50-56. Epub 2017 Dec 4.

Department of Plant & Soil Sciences, University of Kentucky, Lexington, KY, 40546-0091, USA.

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experiments show that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of community composition and the potential interaction between diversity at different scales in maintaining ecosystem functioning remains unclear, especially when many functions are considered (ecosystem multifunctionality). We present an analysis of eight ecosystem functions measured in 65 grasslands worldwide. We find that more diverse grasslands-those with both species-rich local communities (α-diversity) and large compositional differences among localities (β-diversity)-had higher levels of multifunctionality. Moreover, α- and β-diversity synergistically affected multifunctionality, with higher levels of diversity at one scale amplifying the contribution to ecological functions at the other scale. The identity of species influencing ecosystem functioning differed among functions and across local communities, explaining why more diverse grasslands maintained greater functionality when more functions and localities were considered. These results were robust to variation in environmental drivers. Our findings reveal that plant diversity, at both local and landscape scales, contributes to the maintenance of multiple ecosystem services provided by grasslands. Preserving ecosystem functioning therefore requires conservation of biodiversity both within and among ecological communities.
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http://dx.doi.org/10.1038/s41559-017-0395-0DOI Listing
January 2018

Ecological impacts of atmospheric pollution and interactions with climate change in terrestrial ecosystems of the Mediterranean Basin: Current research and future directions.

Environ Pollut 2017 Aug 29;227:194-206. Epub 2017 Apr 29.

Lancaster Environment Center, Lancaster University, Lancaster LA1 4YQ, UK.

Mediterranean Basin ecosystems, their unique biodiversity, and the key services they provide are currently at risk due to air pollution and climate change, yet only a limited number of isolated and geographically-restricted studies have addressed this topic, often with contrasting results. Particularities of air pollution in this region include high O levels due to high air temperatures and solar radiation, the stability of air masses, and dominance of dry over wet nitrogen deposition. Moreover, the unique abiotic and biotic factors (e.g., climate, vegetation type, relevance of Saharan dust inputs) modulating the response of Mediterranean ecosystems at various spatiotemporal scales make it difficult to understand, and thus predict, the consequences of human activities that cause air pollution in the Mediterranean Basin. Therefore, there is an urgent need to implement coordinated research and experimental platforms along with wider environmental monitoring networks in the region. In particular, a robust deposition monitoring network in conjunction with modelling estimates is crucial, possibly including a set of common biomonitors (ideally cryptogams, an important component of the Mediterranean vegetation), to help refine pollutant deposition maps. Additionally, increased attention must be paid to functional diversity measures in future air pollution and climate change studies to establish the necessary link between biodiversity and the provision of ecosystem services in Mediterranean ecosystems. Through a coordinated effort, the Mediterranean scientific community can fill the above-mentioned gaps and reach a greater understanding of the mechanisms underlying the combined effects of air pollution and climate change in the Mediterranean Basin.
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http://dx.doi.org/10.1016/j.envpol.2017.04.062DOI Listing
August 2017

Addition of multiple limiting resources reduces grassland diversity.

Nature 2016 09 24;537(7618):93-96. Epub 2016 Aug 24.

Swiss Federal Institute for Forest, Snow and Landscape Research, Community Ecology, Birmensdorf 8903, Switzerland.

Niche dimensionality provides a general theoretical explanation for biodiversity-more niches, defined by more limiting factors, allow for more ways that species can coexist. Because plant species compete for the same set of limiting resources, theory predicts that addition of a limiting resource eliminates potential trade-offs, reducing the number of species that can coexist. Multiple nutrient limitation of plant production is common and therefore fertilization may reduce diversity by reducing the number or dimensionality of belowground limiting factors. At the same time, nutrient addition, by increasing biomass, should ultimately shift competition from belowground nutrients towards a one-dimensional competitive trade-off for light. Here we show that plant species diversity decreased when a greater number of limiting nutrients were added across 45 grassland sites from a multi-continent experimental network. The number of added nutrients predicted diversity loss, even after controlling for effects of plant biomass, and even where biomass production was not nutrient-limited. We found that elevated resource supply reduced niche dimensionality and diversity and increased both productivity and compositional turnover. Our results point to the importance of understanding dimensionality in ecological systems that are undergoing diversity loss in response to multiple global change factors.
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http://dx.doi.org/10.1038/nature19324DOI Listing
September 2016

Conditional vulnerability of plant diversity to atmospheric nitrogen deposition across the United States.

Proc Natl Acad Sci U S A 2016 Apr 28;113(15):4086-91. Epub 2016 Mar 28.

Department of Botany, University of Wisconsin, Madison, WI 53706.

Atmospheric nitrogen (N) deposition has been shown to decrease plant species richness along regional deposition gradients in Europe and in experimental manipulations. However, the general response of species richness to N deposition across different vegetation types, soil conditions, and climates remains largely unknown even though responses may be contingent on these environmental factors. We assessed the effect of N deposition on herbaceous richness for 15,136 forest, woodland, shrubland, and grassland sites across the continental United States, to address how edaphic and climatic conditions altered vulnerability to this stressor. In our dataset, with N deposition ranging from 1 to 19 kg N⋅ha(-1)⋅y(-1), we found a unimodal relationship; richness increased at low deposition levels and decreased above 8.7 and 13.4 kg N⋅ha(-1)⋅y(-1) in open and closed-canopy vegetation, respectively. N deposition exceeded critical loads for loss of plant species richness in 24% of 15,136 sites examined nationwide. There were negative relationships between species richness and N deposition in 36% of 44 community gradients. Vulnerability to N deposition was consistently higher in more acidic soils whereas the moderating roles of temperature and precipitation varied across scales. We demonstrate here that negative relationships between N deposition and species richness are common, albeit not universal, and that fine-scale processes can moderate vegetation responses to N deposition. Our results highlight the importance of contingent factors when estimating ecosystem vulnerability to N deposition and suggest that N deposition is affecting species richness in forested and nonforested systems across much of the continental United States.
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http://dx.doi.org/10.1073/pnas.1515241113DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4839424PMC
April 2016

How will the semi-natural vegetation of the UK have changed by 2030 given likely changes in nitrogen deposition?

Environ Pollut 2016 Jan 4;208(Pt B):879-89. Epub 2015 Oct 4.

Centre for Ecology & Hydrology, Lancaster Environment Centre, Bailrigg, Lancaster LA1 4AP, UK.

Nitrogen deposition is known to have major impacts on contemporary ecosystems but few studies have addressed how these impacts will develop over coming decades. We consider likely changes to British semi-natural vegetation up to the year 2030 both qualitatively, based on knowledge of species responses from experimental and gradient studies, and quantitatively, based on modelling of species relationships in national monitoring data. We used historical N deposition trends and national predictions of changing deposition to calculate cumulative deposition from 1900 to 2030. Data from the Countryside Survey (1978, 1990 and 1998) was used to parameterise models relating cumulative N deposition to Ellenberg N which were then applied to expected future deposition trends. Changes to habitat suitability for key species of grassland, heathland and bog, and broadleaved woodland to 2030 were predicted using the MultiMOVE model. In UK woodlands by 2030 there is likely to be reduced occurrence of lichens, increased grass cover and a shift towards more nitrophilic vascular plant species. In grasslands we expect changing species composition with reduced occurrence of terricolous lichens and, at least in acid grasslands, reduced species richness. In heaths and bogs we project overall reductions in species richness with decreased occurrence of terricolous lichens and some bryophytes, reduced cover of dwarf shrubs and small increases in grasses. Our study clearly suggests that changes in vegetation due to nitrogen deposition are likely to continue through coming decades.
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http://dx.doi.org/10.1016/j.envpol.2015.09.013DOI Listing
January 2016

Consistent responses of soil microbial communities to elevated nutrient inputs in grasslands across the globe.

Proc Natl Acad Sci U S A 2015 Sep 17;112(35):10967-72. Epub 2015 Aug 17.

Cooperative Institute for Research in Environmental Sciences, University of Colorado, Boulder, CO 80309; Department of Ecology and Evolutionary Biology, University of Colorado, Boulder, CO 80309;

Soil microorganisms are critical to ecosystem functioning and the maintenance of soil fertility. However, despite global increases in the inputs of nitrogen (N) and phosphorus (P) to ecosystems due to human activities, we lack a predictive understanding of how microbial communities respond to elevated nutrient inputs across environmental gradients. Here we used high-throughput sequencing of marker genes to elucidate the responses of soil fungal, archaeal, and bacterial communities using an N and P addition experiment replicated at 25 globally distributed grassland sites. We also sequenced metagenomes from a subset of the sites to determine how the functional attributes of bacterial communities change in response to elevated nutrients. Despite strong compositional differences across sites, microbial communities shifted in a consistent manner with N or P additions, and the magnitude of these shifts was related to the magnitude of plant community responses to nutrient inputs. Mycorrhizal fungi and methanogenic archaea decreased in relative abundance with nutrient additions, as did the relative abundances of oligotrophic bacterial taxa. The metagenomic data provided additional evidence for this shift in bacterial life history strategies because nutrient additions decreased the average genome sizes of the bacterial community members and elicited changes in the relative abundances of representative functional genes. Our results suggest that elevated N and P inputs lead to predictable shifts in the taxonomic and functional traits of soil microbial communities, including increases in the relative abundances of faster-growing, copiotrophic bacterial taxa, with these shifts likely to impact belowground ecosystems worldwide.
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http://dx.doi.org/10.1073/pnas.1508382112DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4568213PMC
September 2015

Plant species' origin predicts dominance and response to nutrient enrichment and herbivores in global grasslands.

Nat Commun 2015 Jul 15;6:7710. Epub 2015 Jul 15.

Department of Entomology, University of Maryland, College Park Maryland 20742, USA.

Exotic species dominate many communities; however the functional significance of species' biogeographic origin remains highly contentious. This debate is fuelled in part by the lack of globally replicated, systematic data assessing the relationship between species provenance, function and response to perturbations. We examined the abundance of native and exotic plant species at 64 grasslands in 13 countries, and at a subset of the sites we experimentally tested native and exotic species responses to two fundamental drivers of invasion, mineral nutrient supplies and vertebrate herbivory. Exotic species are six times more likely to dominate communities than native species. Furthermore, while experimental nutrient addition increases the cover and richness of exotic species, nutrients decrease native diversity and cover. Native and exotic species also differ in their response to vertebrate consumer exclusion. These results suggest that species origin has functional significance, and that eutrophication will lead to increased exotic dominance in grasslands.
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http://dx.doi.org/10.1038/ncomms8710DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4518311PMC
July 2015

Grassland productivity limited by multiple nutrients.

Nat Plants 2015 Jul 6;1:15080. Epub 2015 Jul 6.

Department of Plant and Soil Sciences, University of Kentucky, Lexington, Kentucky 40546, USA.

Terrestrial ecosystem productivity is widely accepted to be nutrient limited(1). Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)(2,3), the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized(4-8). However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+μ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.
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http://dx.doi.org/10.1038/nplants.2015.80DOI Listing
July 2015

Plant diversity predicts beta but not alpha diversity of soil microbes across grasslands worldwide.

Ecol Lett 2015 Jan 28;18(1):85-95. Epub 2014 Nov 28.

CSIRO Land and Water Flagship, Private Bag 5, Wembley, WA, 6913, Australia.

Aboveground-belowground interactions exert critical controls on the composition and function of terrestrial ecosystems, yet the fundamental relationships between plant diversity and soil microbial diversity remain elusive. Theory predicts predominantly positive associations but tests within single sites have shown variable relationships, and associations between plant and microbial diversity across broad spatial scales remain largely unexplored. We compared the diversity of plant, bacterial, archaeal and fungal communities in one hundred and forty-five 1 m(2) plots across 25 temperate grassland sites from four continents. Across sites, the plant alpha diversity patterns were poorly related to those observed for any soil microbial group. However, plant beta diversity (compositional dissimilarity between sites) was significantly correlated with the beta diversity of bacterial and fungal communities, even after controlling for environmental factors. Thus, across a global range of temperate grasslands, plant diversity can predict patterns in the composition of soil microbial communities, but not patterns in alpha diversity.
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http://dx.doi.org/10.1111/ele.12381DOI Listing
January 2015

Soil phosphorus constrains biodiversity across European grasslands.

Glob Chang Biol 2014 Dec 21;20(12):3814-22. Epub 2014 Jul 21.

Plant Conservation and Population Biology, Department Biology, University of Leuven, Kasteelpark Arenberg 31, Leuven, B-3001, Belgium.

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. On the basis of 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 toward 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.
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http://dx.doi.org/10.1111/gcb.12650DOI Listing
December 2014

Herbivores and nutrients control grassland plant diversity via light limitation.

Nature 2014 Apr 9;508(7497):517-20. Epub 2014 Mar 9.

Instituto de Investigaciones Marinas y Costeras (IIMyC), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Mar del Plata 7600 , Argentina.

Human alterations to nutrient cycles and herbivore communities are affecting global biodiversity dramatically. 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 systems. 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.
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http://dx.doi.org/10.1038/nature13144DOI Listing
April 2014

Eutrophication weakens stabilizing effects of diversity in natural grasslands.

Nature 2014 Apr 16;508(7497):521-5. Epub 2014 Feb 16.

Swiss Federal Institute for Forest, Snow and Landscape Research, 8903 Birmensdorf, Switzerland.

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.
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http://dx.doi.org/10.1038/nature13014DOI Listing
April 2014

Personal nitrogen footprint tool for the United Kingdom.

Environ Sci Process Impacts 2014 Jul;16(7):1563-9

Lancaster Environment Centre, Lancaster University, Lancaster, LA1 4YQ, UK.

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.
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http://dx.doi.org/10.1039/c3em00690eDOI Listing
July 2014

Soil chemistry and fertility alterations in response to N application in a semiarid Mediterranean shrubland.

Sci Total Environ 2013 May 15;452-453:78-86. Epub 2013 Mar 15.

Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas (MNCN-CSIC), Madrid, Spain.

N deposition is currently affecting nutrient cycling in terrestrial ecosystems. We studied the effects of four years of N application (0, 10, 20 and 50 kg N ha(-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.
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http://dx.doi.org/10.1016/j.scitotenv.2013.02.049DOI Listing
May 2013