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Land‐use intensification has caused dramatic declines in grassland biodiversity, and in Europe this decline has not yet been halted. Therefore, a major challenge in ecological restoration is to effectively restore plant diversity in permanent grasslands, that is, long‐term managed grassland not being used as cropland. As species richness is often limited by seed dispersal, introducing locally absent species is crucial. Seeding diverse seed mixtures is an appropriate tool, but restoration outcomes are highly variable. Considering constraints by land‐use intensity and plant functional traits during restoration could improve the efficacy of restoration efforts.We established a full‐factorial experiment with high‐diversity seeding and topsoil disturbance in 73 grasslands along land‐use intensity and productivity gradients and evaluated the restoration success over 5 years. In addition, after 5 years we assessed the number of established species and tested whether specific traits promoted the establishment of sown species in interaction with productivity.Plant diversity increased considerably when seeding and disturbance were combined while seeding alone only slightly increased species richness. Along land‐use gradients, we found no direct effects of grazing or mowing on the establishment of sown species, but fertilization indirectly limited establishment in the combined treatment via a negative effect of productivity.Functional traits structured the establishment of sown species, especially in the combined treatments, but effects varied with productivity. Species with low seed mass and low specific leaf area had greater establishment success in low‐productivity grasslands, whereas species with high seed mass had greater establishment in high‐productivity grasslands.Synthesis and applications. Our results highlight the potential of using diverse seed mixtures for biodiversity restoration in a wide range of permanent grasslands, especially when combined with topsoil disturbance. However, productivity both limited restoration success and affected establishment via an interaction with plant functional traits. As productivity is a major constraint in grassland restoration, we strongly recommend to first stop fertilization and reduce soil nutrient loads in highly productive grasslands. Restoration by seeding, with or without disturbance, should only be applied when productivity is reduced and the local environment should be considered to compile site‐specific seed mixtures to maximize restoration success. ​ [ABSTRACT FROM AUTHOR]

Abstract: 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. [ABSTRACT FROM AUTHOR]

Soil seed banks support the recovery of plant species and communities after disturbances. Against the background of increasing disturbance frequencies due to current changes in climate and land use, the importance of seed banks to maintain ecosystem functioning of grasslands is likely to increase. This makes seed bank ecology a highly relevant topic of studies on ecosystem functioning. One of the main drivers of seed bank composition and density in grasslands is the current management. Although highly intensive grassland management can affect soil seed banks negatively, it is still unclear which component of management – fertilization, mowing or grazing – is the most influential. We studied soil seed banks of 73 mesic grasslands, which were arranged along a gradient of land-use intensity, by two seasonally differing surveys. Grasslands generally exhibited rather poor seed banks with low numbers of species and seedlings. Both were even lower in spring compared to autumn sampling. Grassland management turned out to affect soil seed banks in a complex way, via direct and indirect paths. While intensive mowing and fertilization decreased species richness and seedling density, grazing intensity increased species richness in the seed bank. Thus, land-use intensity, an index calculated as a sum of all three components, had highly ambivalent effects on seed banks. We conclude that the positive effect of grazing on soil seed banks makes pastures more resilient towards mechanic disturbances as compared to meadows. [ABSTRACT FROM AUTHOR]

Plant functional traits reflect individual and community ecological strategies. They allow the detection of directional changes in community dynamics and ecosystemic processes, being an additional tool to assess biodiversity than species richness. Analysis of functional patterns in plant communities provides mechanistic insight into biodiversity alterations due to anthropogenic activity. Although studies have consi-dered of either anthropogenic management or nutrient availability on functional traits in temperate grasslands, studies combining effects of both drivers are scarce. Here, we assessed the impacts of management intensity (fertilization, mowing, grazing), nutrient stoichiometry (C, N, P, K), and vegetation composition on community-weighted means ( CWMs) and functional diversity (Rao's Q) from seven plant traits in 150 grasslands in three regions in Germany, using data of 6 years. Land use and nutrient stoichiometry accounted for larger proportions of model variance of CWM and Rao's Q than species richness and productivity. Grazing affected all analyzed trait groups; fertilization and mowing only impacted generative traits. Grazing was clearly associated with nutrient retention strategies, that is, investing in durable structures and production of fewer, less variable seed. Phenological variability was increased. Fertilization and mowing decreased seed number/mass variability, indicating competition-related effects. Impacts of nutrient stoichiometry on trait syndromes varied. Nutrient limitation (large N:P, C:N ratios) promoted species with conservative strategies, that is, investment in durable plant structures rather than fast growth, fewer seed, and delayed flowering onset. In contrast to seed mass, leaf-economics variability was reduced under P shortage. Species diversity was positively associated with the variability of generative traits. Synthesis. Here, land use, nutrient availability, species richness, and plant functional strategies have been shown to interact complexly, driving community composition, and vegetation responses to management intensity. We suggest that deeper understanding of underlying mechanisms shaping community assembly and biodiversity will require analyzing all these parameters. [ABSTRACT FROM AUTHOR]

Aims The relationship between biodiversity and ecosystem functioning is among the most active areas of ecological research. Furthermore, enhancing the diversity of degraded ecosystems is a major goal in applied restoration ecology. In grasslands, many species may be locally absent due to dispersal or microsite limitation and may therefore profit from mechanical disturbance of the resident vegetation. We established a seed addition and disturbance experiment across several grassland sites of different land use to test whether plant diversity can be increased in these grasslands. Additionally, the experiment will allow us testing the consequences of increased plant diversity for ecosystem processes and for the diversity of other taxa in real-world ecosystems. Here, we present details of the experimental design and report results from the first vegetation survey 1 year after disturbance and seed addition. Moreover, we tested whether the effects of seed addition and disturbance varied among grassland depending on their land use or pre-disturbance plant diversity. Methods A full-factorial experiment was installed in 73 grasslands in three regions across Germany. Grasslands were under regular agricultural use, but varied in the type and the intensity of management, thereby representing the range of management typical for large parts of Central Europe. The disturbance treatment consisted of disturbing the top 10 cm of the sward using a rotavator or rotary harrow. Seed addition consisted of sowing a high-diversity seed mixture of regional plant species. These species were all regionally present, but often locally absent, depending on the resident vegetation composition and richness of each grassland. Important Findings Seven months after sward disturbance, respective plots had significantly increased in bare soil, seedling species richness and number of seedlings. Seed addition had increased plant species richness, but only in combination with sward disturbance. The increase in species richness, when both seed addition and disturbance was applied, was higher at high land-use intensity and low resident diversity. Thus, we show that at least the early recruitment of many species is possible also at high land-use intensity, indicating the potential to restore and enhance biodiversity of species-poor agricultural grasslands. Our newly established experiment provides a unique platform for broad-scale research on the land-use dependence of future trajectories of vegetation diversity and composition and their effects on ecosystem functioning. [ABSTRACT FROM AUTHOR]

Land-use intensification, nutrient enrichment and enhanced productivity levels are major drivers of the current biodiversity loss in agricultural grasslands. Although these drivers are potentially strongly causally linked, their potential to explain changes in biodiversity can enhance each other and often varies over time. Here, we explored the explanatory power of different abiotic drivers and the Ellenberg indicator value for nutrient availability to predict plant species richness using data of 150 grassland plots over seven years. The approach also allows to assess the temporal robustness of many of these drivers. Our results revealed the Ellenberg indicator value for nutrients to be the best and temporally most consistent predictor of plant species richness. Unweighted mean Ellenberg values showed an even stronger negative relationship to plant species richness than abundance weighted mean values. Direct measurements of soil and plant biomass nutrients, aboveground biomass and land-use intensity predicted plant species richness less well and showed considerable temporal variability in the respective association. In case of biomass nitrogen and phosphorus concentrations, current year weather conditions are assumed to strongly affect the strength of the relationship by shifting the phenological development considerable among sampling years. Although mainly driven by aboveground biomass, a combination of biomass nutrient concentrations with productivity, i.e. the nutrient stock in the aboveground biomass, turned out to better predict plant species richness than its single components and inter-annual variability in the relationship was markedly lower. Especially for aboveground biomass and the intensity of land-use, long-term mean values significantly improved the explanatory power compared to single year measurements. We conclude that studies explaining biodiversity changes should carefully take into account temporal variability in the drivers under study before judging the strength of a particular mechanistic relationship. [ABSTRACT FROM AUTHOR]