Nitrogen availability and plant functional composition modify biodiversity-multifunctionality relationships.

Biodiversity typically increases multiple ecosystem functions simultaneously (multifunctionality) but variation in the strength and direction of biodiversity effects between studies suggests context dependency. To determine how different factors modulate the diversity effect on multifunctionality, we established a large grassland experiment manipulating plant species richness, resource addition, functional composition (exploitative vs. conservative species), functional diversity and enemy abundance. We measured ten above- and belowground functions and calculated ecosystem multifunctionality. Species richness and functional diversity both increased multifunctionality, but their effects were context dependent. Richness increased multifunctionality when communities were assembled with fast-growing species. This was because slow species were more redundant in their functional effects, whereas different fast species promoted different functions. Functional diversity also increased multifunctionality but this effect was dampened by nitrogen enrichment and enemy presence. Our study suggests that a shift towards fast-growing communities will not only alter ecosystem functioning but also the strength of biodiversity-functioning relationships.

Drought effects on trait space of winter wheat are independent of land management.

Investigating plant responses to climate change is key to develop suitable adaptation strategies. However, whether changes in land management can alleviate increasing drought threats to crops in the future is still unclear. We conducted a management × drought experiment with winter wheat (Triticum aestivum L.) to study plant water and vegetative traits in response to drought and management (conventional vs organic farming, with intensive vs conservation tillage). Water traits (root water uptake pattern, stem metaxylem area, leaf water potential, stomatal conductance) and vegetative traits (plant height, leaf area, leaf Chl content) were considered simultaneously to characterise the variability of multiple traits in a trait space, using principal component analysis. Management could not alleviate the drought impacts on plant water traits as it mainly affected vegetative traits, with yields ultimately being affected by both management and drought. Trait spaces were clearly separated between organic and conventional management as well as between drought and control conditions. Moreover, changes in trait space triggered by management and drought were independent from each other. Neither organic management nor conservation tillage eased drought impacts on winter wheat. Thus, our study raised concerns about the effectiveness of these management options as adaptation strategies to climate change.

Land-use intensity alters networks between biodiversity, ecosystem functions, and services.

Land-use intensification can increase provisioning ecosystem services, such as food and timber production, but it also drives changes in ecosystem functioning and biodiversity loss, which may ultimately compromise human wellbeing. To understand how changes in land-use intensity affect the relationships between biodiversity, ecosystem functions, and services, we built networks from correlations between the species richness of 16 trophic groups, 10 ecosystem functions, and 15 ecosystem services. We evaluated how the properties of these networks varied across land-use intensity gradients for 150 forests and 150 grasslands. Land-use intensity significantly affected network structure in both habitats. Changes in connectance were larger in forests, while changes in modularity and evenness were more evident in grasslands. Our results show that increasing land-use intensity leads to more homogeneous networks with less integration within modules in both habitats, driven by the belowground compartment in grasslands, while forest responses to land management were more complex. Land-use intensity strongly altered hub identity and module composition in both habitats, showing that the positive correlations of provisioning services with biodiversity and ecosystem functions found at low land-use intensity levels, decline at higher intensity levels. Our approach provides a comprehensive view of the relationships between multiple components of biodiversity, ecosystem functions, and ecosystem services and how they respond to land use. This can be used to identify overall changes in the ecosystem, to derive mechanistic hypotheses, and it can be readily applied to further global change drivers.

Additive effects of two agri-environmental schemes on plant diversity but not on productivity indicators in permanent grasslands in Switzerland.

Different agri-environmental schemes (AES), such as ecological focus areas and organic farming, have been suggested to reduce the impact of intensive agriculture on the environment and to conserve or even restore farmland biodiversity. However, the effectiveness of such schemes, their ability to actually support biodiversity and associated trade-offs with agricultural production are still debated. We analysed a large dataset from the biodiversity monitoring in the Swiss agricultural landscape to assess the effects of two different grassland AES, i.e., extensively managed ecological focus areas (EFAs versus non-EFAs) and organic farming (versus conventional), on plant diversity, plant community composition and productivity indicators, i.e., weed abundance, forage value and nutrient availability. We also considered environmental factors, i.e., topography and soil conditions, which potentially modulate AES effects on biodiversity. We used in total 1170 plots in permanent grasslands, managed as meadows or pastures. Both AES had significant positive effects on plant diversity. However, EFAs increased plant richness considerably stronger (+6.6 species) than organic farming (+1.8 species). Effects of the two schemes were additive with organic EFA grasslands exhibiting highest plant diversity. Differences in topography partly explained AES effects on diversity as both AES were associated with differences in elevation and slope. Thus, future assessments of the effectiveness of AES need to consider the non-random placement of AES across heterogeneous landscapes. EFA grasslands revealed a considerably reduced agricultural productivity as shown by low forage values and low nutrient availability. Yet, the abundance of agricultural weeds, i.e., agriculturally undesired plant species, was lower in EFA compared to non-EFA grasslands. Productivity indicators were only weakly affected by organic farming and other than for plant diversity, productivity did not differ between organic and conventional EFA grasslands. The positive additive diversity effects of EFAs and organic grassland farming underline the potential of both AES to contribute to biodiversity conservation in agricultural landscapes, though to a different extent. Comparing the effects of the two AES revealed that the lower the reduction in agricultural productivity associated with an AES, the smaller the gains in plant diversity, highlighting the inevitable trade-off between productivity and plant diversity in semi-natural grasslands.

Microbial drivers of plant richness and productivity in a grassland restoration experiment along a gradient of land-use intensity.

Plant-soil feedbacks (PSFs) underlying grassland plant richness and productivity are typically coupled with nutrient availability; however, we lack understanding of how restoration measures to increase plant diversity might affect PSFs. We examined the roles of sward disturbance, seed addition and land-use intensity (LUI) on PSFs. We conducted a disturbance and seed addition experiment in 10 grasslands along a LUI gradient and characterized plant biomass and richness, soil microbial biomass, community composition and enzyme activities. Greater plant biomass at high LUI was related to a decrease in the fungal to bacterial ratios, indicating highly productive grasslands to be dominated by bacteria. Lower enzyme activity per microbial biomass at high plant species richness indicated a slower carbon (C) cycling. The relative abundance of fungal saprotrophs decreased, while pathogens increased with LUI and disturbance. Both fungal guilds were negatively associated with plant richness, indicating the mechanisms underlying PSFs depended on LUI. We show that LUI and disturbance affect fungal functional composition, which may feedback on plant species richness by impeding the establishment of pathogen-sensitive species. Therefore, we highlight the need to integrate LUI including its effects on PSFs when planning for practices that aim to optimize plant diversity and productivity.

Biodiversity at multiple trophic levels is needed for ecosystem multifunctionality.

Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.

Cropping systems alter hydraulic traits of barley but not pea grown in mixture.

Extreme events such as drought and heatwaves are among the biggest challenges to agricultural production and food security. However, the effects of cropping systems on drought resistance of arable crops via their hydraulic behaviour remain unclear. We investigated how hydraulic traits of a field-grown pea-barley (Pisum sativum L. and Hordeum vulgare L.) mixture were affected by different cropping systems, that is, organic and conventional farming with intensive or conservation tillage. Xylem vulnerability to cavitation of both species was estimated by measuring the pressure inducing 50% loss of hydraulic conductivity (P50 ), while the water stress plants experienced in the field were assessed using native percentage loss of hydraulic conductivity (nPLC). Pea and barley showed contrasting hydraulic behaviours: pea was less vulnerable to xylem cavitation and less stressed than barley; cropping systems affected the xylem vulnerability of barley, but not of pea. Barley grown under conventional farming with no tillage was more vulnerable and stressed than under organic farming with intensive tillage. nPLC proved to be a valuable indicator for plant water stress. Our results highlight the impact of cropping systems on crop xylem vulnerability and drought resistance, thus plant hydraulic traits, for protecting food security under future climate.

Land use intensity, rather than plant species richness, affects the leaching risk of multiple nutrients from permanent grasslands.

The intensification of land use constitutes one of the main drivers of global change and alters nutrient fluxes on all spatial scales, causing landscape-level eutrophication and contamination of natural resources. Changes in soil nutrient concentrations are thus indicative for crucial environmental issues associated with intensive land use. We measured concentrations of NO3 -N, NH4 -N, P, K, Mg, and Ca using 1,326 ion-exchange resin bags buried in 20 cm depth beneath the main root zone in 150 temperate grasslands. Nutrient concentrations were related to land use intensity, that is, fertilization, mowing, grazing intensities, and plant diversity by structural equation modeling. Furthermore, we assessed the response of soil nutrients to mechanical sward disturbance and subsequent reseeding, a common practice for grassland renewal. Land use intensity, especially fertilization, significantly increased the concentrations of NO3 -N, NH4 -N, K, P, and also Mg. Besides fertilization (and tightly correlated mowing) intensity, grazing strongly increased NO3 -N and K concentrations. Plant species richness decreased P and NO3 -N concentrations in soil when grassland productivity of the actual year was statistically taken into account, but not when long-term averages of productivity were used. Thus, we assume that, in the actual study year, a distinct drought period might have caused the observed decoupling of productivity from fertilization and soil nutrients. Breaking up the grassland sward drastically increased NO3 -N concentrations (+146%) but reduced NH4 -N, P, and K concentrations, unbalancing soil nutrient stoichiometry and boosting the risk of N leaching. Reseeding the sward after disturbance did not have a short-term effect on nutrient concentrations. We conclude that renewal of permanent grassland should be avoided as far as possible and future grassland management has to strongly rise the effectiveness of fertilization. Additionally, grassland management might have to increasingly taking care of periods of drought, in which nutrient additions might not increase plant growth but potentially only facilitate leaching.

Interannual variation in land-use intensity enhances grassland multidiversity.

Although temporal heterogeneity is a well-accepted driver of biodiversity, effects of interannual variation in land-use intensity (LUI) have not been addressed yet. Additionally, responses to land use can differ greatly among different organisms; therefore, overall effects of land-use on total local biodiversity are hardly known. To test for effects of LUI (quantified as the combined intensity of fertilization, grazing, and mowing) and interannual variation in LUI (SD in LUI across time), we introduce a unique measure of whole-ecosystem biodiversity, multidiversity. This synthesizes individual diversity measures across up to 49 taxonomic groups of plants, animals, fungi, and bacteria from 150 grasslands. Multidiversity declined with increasing LUI among grasslands, particularly for rarer species and aboveground organisms, whereas common species and belowground groups were less sensitive. However, a high level of interannual variation in LUI increased overall multidiversity at low LUI and was even more beneficial for rarer species because it slowed the rate at which the multidiversity of rare species declined with increasing LUI. In more intensively managed grasslands, the diversity of rarer species was, on average, 18% of the maximum diversity across all grasslands when LUI was static over time but increased to 31% of the maximum when LUI changed maximally over time. In addition to decreasing overall LUI, we suggest varying LUI across years as a complementary strategy to promote biodiversity conservation.

Land use intensification alters ecosystem multifunctionality via loss of biodiversity and changes to functional composition.

Global change, especially land-use intensification, affects human well-being by impacting the delivery of multiple ecosystem services (multifunctionality). However, whether biodiversity loss is a major component of global change effects on multifunctionality in real-world ecosystems, as in experimental ones, remains unclear. Therefore, we assessed biodiversity, functional composition and 14 ecosystem services on 150 agricultural grasslands differing in land-use intensity. We also introduce five multifunctionality measures in which ecosystem services were weighted according to realistic land-use objectives. We found that indirect land-use effects, i.e. those mediated by biodiversity loss and by changes to functional composition, were as strong as direct effects on average. Their strength varied with land-use objectives and regional context. Biodiversity loss explained indirect effects in a region of intermediate productivity and was most damaging when land-use objectives favoured supporting and cultural services. In contrast, functional composition shifts, towards fast-growing plant species, strongly increased provisioning services in more inherently unproductive grasslands.

Effects of management practices on the ecosystem-service multifunctionality of temperate grasslands.

Human wellbeing depends on ecosystem services, highlighting the need for improving the ecosystem-service multifunctionality of food and feed production systems. We study Swiss agricultural grasslands to assess how employing and combining three widespread aspects of grassland management and their interactions can enhance 22 plot-level ecosystem service indicators, as well as ecosystem-service multifunctionality. The three management aspects we assess are i) organic production system, ii) an eco-scheme prescribing extensive management (without fertilization), and iii) harvest type (pasture vs. meadow). While organic production system and interactions between the three management aspects play a minor role, the main effects of eco-scheme and harvest type considerably shape single services. Moreover, the eco-scheme 'extensive management' and the harvest type 'pasture' enhance plot-scale ecosystem-service multifunctionality, mostly through facilitating cultural services at the expense of provisioning services. These changes in ecosystem-service supply occur mainly via changes in land-use intensity, i.e., reduced fertilizer input and harvest frequency. In conclusion, diversifying grassland management where this is currently homogeneous across farms and landscapes depicts an important first step to improve landscape-scale multifunctionality for sustainable grassland systems. To meet societal ecosystem services demand, the three studied management aspects can be systematically combined to increase ecosystem services that are in short supply.

Convergence and divergence in science and practice of urban and rural forest restoration.

Forest restoration has never been higher on policymakers' agendas. Complex and multi-dimensional arrangements across the urban-rural continuum challenge restorationists and require integrative approaches to strengthen environmental protection and increase restoration outcomes. It remains unclear if urban and rural forest restoration are moving towards or away from each other in practice and research, and whether comparing research outcomes can help stakeholders to gain a clearer understanding of the interconnectedness between the two fields. This study aims to identify the challenges and opportunities for enhancing forest restoration in both urban and rural systems by reviewing the scientific evidence, engaging with key stakeholders and using an urban-rural forest restoration framework. Using the Society for Ecological Restoration's International Principles as discussion topics, we highlight aspects of convergence and divergence between the two fields to broaden our understanding of forest restoration and promote integrative management approaches to address future forest conditions. Our findings reveal that urban and rural forest restoration have convergent and divergent aspects. We emphasise the importance of tailoring goals and objectives to specific contexts and the need to design different institutions and incentives based on the social and ecological needs and goals of stakeholders in different regions. Additionally, we discuss the challenges of achieving high levels of ecological restoration and the need to go beyond traditional ecology to plan, implement, monitor, and adaptively manage restored forests. We suggest that rivers and watersheds could serve as a common ground linking rural and urban landscapes and that forest restoration could interact with other environmental protection measures. We note the potential for expanding the creative vision associated with increasing tree-containing environments in cities to generate more diverse and resilient forest restoration outcomes in rural settings. This study underscores the value of integrative management approaches in addressing future forest conditions across the urban-rural continuum. Our framework provides valuable insights for policymakers, researchers, and decision-makers to advance the field of forest restoration and address the challenges of restoration across the urban-rural continuum. The rural-urban interface serves as a convergence point for forest restoration, and both urban and rural fields can benefit from each other's expertise.

A slow-fast trait continuum at the whole community level in relation to land-use intensification.

Organismal functional strategies form a continuum from slow- to fast-growing organisms, in response to common drivers such as resource availability and disturbance. However, whether there is synchronisation of these strategies at the entire community level is unclear. Here, we combine trait data for >2800 above- and belowground taxa from 14 trophic guilds spanning a disturbance and resource availability gradient in German grasslands. The results indicate that most guilds consistently respond to these drivers through both direct and trophically mediated effects, resulting in a 'slow-fast' axis at the level of the entire community. Using 15 indicators of carbon and nutrient fluxes, biomass production and decomposition, we also show that fast trait communities are associated with faster rates of ecosystem functioning. These findings demonstrate that 'slow' and 'fast' strategies can be manifested at the level of whole communities, opening new avenues of ecosystem-level functional classification.

The supply of multiple ecosystem services requires biodiversity across spatial scales.

The impact of local biodiversity loss on ecosystem functioning is well established, but the role of larger-scale biodiversity dynamics in the delivery of ecosystem services remains poorly understood. Here we address this gap using a comprehensive dataset describing the supply of 16 cultural, regulating and provisioning ecosystem services in 150 European agricultural grassland plots, and detailed multi-scale data on land use and plant diversity. After controlling for land-use and abiotic factors, we show that both plot-level and surrounding plant diversity play an important role in the supply of cultural and aboveground regulating ecosystem services. In contrast, provisioning and belowground regulating ecosystem services are more strongly driven by field-level management and abiotic factors. Structural equation models revealed that surrounding plant diversity promotes ecosystem services both directly, probably by fostering the spill-over of ecosystem service providers from surrounding areas, and indirectly, by maintaining plot-level diversity. By influencing the ecosystem services that local stakeholders prioritized, biodiversity at different scales was also shown to positively influence a wide range of stakeholder groups. These results provide a comprehensive picture of which ecosystem services rely most strongly on biodiversity, and the respective scales of biodiversity that drive these services. This key information is required for the upscaling of biodiversity-ecosystem service relationships, and the informed management of biodiversity within agricultural landscapes.

Enzyme kinetics inform about mechanistic changes in tea litter decomposition across gradients in land-use intensity in Central German grasslands.

Grassland ecosystems provide important ecosystem services such as nutrient cycling and primary production that are affected by land-use intensity. To assess the effects of land-use intensity, operational and sensitive ecological indicators that integrate effects of grassland management on ecosystem processes such as organic matter turnover are needed. Here, we investigated the suitability of measuring the mass loss of standardized tea litter together with extracellular enzyme kinetics as a proxy of litter decomposition in the topsoil of grasslands along a well-defined land-use intensity gradient (fertilization, mowing, grazing) in Central Germany. Tea bags containing either green tea (high-quality litter) or rooibos tea (low-quality litter) were buried in 5 cm soil depth. Litter mass loss was measured after three (early-stage decomposition) and 12 months (mid-stage decomposition). Based on the fluorescence measurement of the reaction product 4-methylumbelliferone, Michaelis-Menten enzyme kinetics (Vmax: potential maximum rate of activity; Km: substrate affinity) of five hydrolases involved in the carbon (C)-, nitrogen (N)- and phosphorus (P)-cycle (ß-glucosidase (BG), cellobiohydrolase (CBH), cellotriohydrolase (CTH), 1,4-ß-N-acetylglucosaminidase (NAG), and phosphatase (PH)) were determined in tea litter bags and in the surrounding soil. The land-use intensity index (LUI), summarizing fertilization, mowing, grazing, and in particular the frequency of mowing were identified as important drivers of early-stage tea litter decomposition. Mid-stage decomposition was influenced by grazing intensity. The higher the potential activity of all measured C-, N- and P-targeting enzymes, the higher was the decomposition of both tea litters in the early-phase. During mid-stage decomposition, individual enzyme parameters (Vmax of CTH and PH, Km of CBH) became more important. The tea bag method proved to be a suitable indicator which allows an easy and cost-effective assessment of land-use intensity effects on decay processes in manged grasslands. In combination with enzyme kinetics it is an appealing approach to identify mechanisms driving litter break down.

The Evolution of Ecological Diversity in Acidobacteria.

Acidobacteria occur in a large variety of ecosystems worldwide and are particularly abundant and highly diverse in soils. In spite of their diversity, only few species have been characterized to date which makes Acidobacteria one of the most poorly understood phyla among the domain Bacteria. We used a culture-independent niche modeling approach to elucidate ecological adaptations and their evolution for 4,154 operational taxonomic units (OTUs) of Acidobacteria across 150 different, comprehensively characterized grassland soils in Germany. Using the relative abundances of their 16S rRNA gene transcripts, the responses of active OTUs along gradients of 41 environmental variables were modeled using hierarchical logistic regression (HOF), which allowed to determine values for optimum activity for each variable (niche optima). By linking 16S rRNA transcripts to the phylogeny of full 16S rRNA gene sequences, we could trace the evolution of the different ecological adaptations during the diversification of Acidobacteria. This approach revealed a pronounced ecological diversification even among acidobacterial sister clades. Although the evolution of habitat adaptation was mainly cladogenic, it was disrupted by recurrent events of convergent evolution that resulted in frequent habitat switching within individual clades. Our findings indicate that the high diversity of soil acidobacterial communities is largely sustained by differential habitat adaptation even at the level of closely related species. A comparison of niche optima of individual OTUs with the phenotypic properties of their cultivated representatives showed that our niche modeling approach (1) correctly predicts those physiological properties that have been determined for cultivated species of Acidobacteria but (2) also provides ample information on ecological adaptations that cannot be inferred from standard taxonomic descriptions of bacterial isolates. These novel information on specific adaptations of not-yet-cultivated Acidobacteria can therefore guide future cultivation trials and likely will increase their cultivation success.

Changes in plant-herbivore network structure and robustness along land-use intensity gradients in grasslands and forests.

Land-use intensification poses major threats to biodiversity, such as to insect herbivore communities. The stability of these communities depends on interactions linking herbivores and host plants. How interaction network structure begets robustness, and thus stability, in different ecosystems and how network structure and robustness are altered along land-use intensity gradients are unclear. We analyzed plant-herbivore networks based on literature-derived interactions and long-term sampling from 289 grasslands and forests in three regions of Germany. Network size and nestedness were the most important determinants of network robustness in both ecosystems. Along land-use intensity gradients, networks in moderately grazed grasslands were more robust than in those managed by frequent mowing or fertilization. In forests, changes of network robustness along land-use intensity gradients relied on changes in plant species richness. Our results expand our knowledge of the stability of plant-herbivore networks and indicate options for management aimed at stabilizing herbivore communities.

Above- and belowground biodiversity jointly tighten the P cycle in agricultural grasslands.

Experiments showed that biodiversity increases grassland productivity and nutrient exploitation, potentially reducing fertiliser needs. Enhancing biodiversity could improve P-use efficiency of grasslands, which is beneficial given that rock-derived P fertilisers are expected to become scarce in the future. Here, we show in a biodiversity experiment that more diverse plant communities were able to exploit P resources more completely than less diverse ones. In the agricultural grasslands that we studied, management effects either overruled or modified the driving role of plant diversity observed in the biodiversity experiment. Nevertheless, we show that greater above- (plants) and belowground (mycorrhizal fungi) biodiversity contributed to tightening the P cycle in agricultural grasslands, as reduced management intensity and the associated increased biodiversity fostered the exploitation of P resources. Our results demonstrate that promoting a high above- and belowground biodiversity has ecological (biodiversity protection) and economical (fertiliser savings) benefits. Such win-win situations for farmers and biodiversity are crucial to convince farmers of the benefits of biodiversity and thus counteract global biodiversity loss.

Contrasting responses of above- and belowground diversity to multiple components of land-use intensity.

Land-use intensification is a major driver of biodiversity loss. However, understanding how different components of land use drive biodiversity loss requires the investigation of multiple trophic levels across spatial scales. Using data from 150 agricultural grasslands in central Europe, we assess the influence of multiple components of local- and landscape-level land use on more than 4,000 above- and belowground taxa, spanning 20 trophic groups. Plot-level land-use intensity is strongly and negatively associated with aboveground trophic groups, but positively or not associated with belowground trophic groups. Meanwhile, both above- and belowground trophic groups respond to landscape-level land use, but to different drivers: aboveground diversity of grasslands is promoted by diverse surrounding land-cover, while belowground diversity is positively related to a high permanent forest cover in the surrounding landscape. These results highlight a role of landscape-level land use in shaping belowground communities, and suggest that revised agroecosystem management strategies are needed to conserve whole-ecosystem biodiversity.

Drought boosts risk of nitrate leaching from grassland fertilisation.

Both climate change and agricultural intensification are drivers of global nutrient cycles and biodiversity loss. A potentially great environmental threat can arise when these two drivers interact, for example, when farmers try to compensate reduced soil nutrient availability due to drought by the application of liquid organic fertiliser. As dry soils don't hold back nutrients very well, this approach can lead to nitrate leaching and potentially also to the pollution of drinking water. However, little is known about leaching from dry but fertilised grassland soil, and how this is affected by land use intensity and plant diversity. In this mesocosm study, we transferred 60 grassland sods differing in past land use intensity to a greenhouse and treated them with severe drought, fertilisation and both together. Drought was induced by almost entirely stopping irrigation for seven weeks. Fertilisation was done by three applications of slurry summing up to 168 kg total nitrogen per hectare (111 kg NH4-N). We assessed nutrient leaching risk with ion-exchange resin (IER) bags installed in the soil of all mesocosms. IER bags were retrieved after drought and extracts were analysed for concentrations of nitrate, ammonium, phosphate and potassium. Fertilisation partially buffered drought-induced losses in yield. However, the interaction of fertilisation and drought resulted in a drastic increase in nitrate leaching risk when soils are rewetted (>300%), while neither drought nor fertilisation alone were significant. Ammonium concentrations followed the same trend as nitrate, but less pronounced. Phosphate and potassium concentrations were not affected by the treatments. Past land use was hardly related to soil nutrient concentrations, rather was plant diversity. However, results indicate that plant diversity was not driving nitrate and ammonium concentrations under drought and/or fertilisation. This study reveals grassland fertilisation during drought to be a severe environmental problem due to significantly increased nitrate leaching risk.