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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. Sustainable agricultural policies need to be practically assessed. Here, the authors assess how management practices affect ecosystem services in Swiss agricultural grasslands showing that organic farming has a lesser impact than the eco-scheme and the use as pasture or meadow.

Human land use may detrimentally affect biodiversity, yet long-term stability of species communities is vital for maintaining ecosystem functioning. Community stability can be achieved by higher species diversity (portfolio effect), higher asynchrony across species (insurance hypothesis) and higher abundance of populations. However, the relative importance of these stabilizing pathways and whether they interact with land use in real-world ecosystems is unknown. We monitored inter-annual fluctuations of 2,671 plant, arthropod, bird and bat species in 300 sites from three regions. Arthropods show 2.0-fold and birds 3.7-fold higher community fluctuations in grasslands than in forests, suggesting a negative impact of forest conversion. Land-use intensity in forests has a negative net impact on stability of bats and in grasslands on birds. Our findings demonstrate that asynchrony across species—much more than species diversity alone—is the main driver of variation in stability across sites and requires more attention in sustainable management. Long-term stability of ecological communities is vital for maintaining ecosystem functioning. Here, Blüthgen et al . show that greater land-use intensity in grasslands and forests can have negative impacts on the stability of plant and animal communities, driven primarily by variation in asynchrony between species.

Globally, we face a dramatic biodiversity loss in agricultural systems as well as severe ecosystem degradation. In grasslands, higher biodiversity in terms of plant diversity was shown to increase the diversity of higher trophic levels and provide benefits for farmers such as higher and more stable yields. However, we lack a systematic overview of costs for more diverse seed mixtures, which are an essential tool in maintaining and increasing plant diversity in grasslands. We here investigated the prices and characteristics of 262 commercially available seed mixtures from six German or Swiss online shops and quantified the relationships between seed mixture prices and plant diversity. The most frequent seed mixtures contained 1–10 species and were designed for rather intensive grassland management. On the contrary, a smaller set of seed mixtures with particularly high plant diversity (>30 species), usually of native ecotypes, were offered for restoration purposes. More diverse seed mixtures were also more expensive. For example, a seed mixture with 10 or 30 species was on average +63% or +387% more expensive, respectively, than a product containing only one species. The relationship between plant diversity and seed mixture prices per ha was related to other seed mixture characteristics, of which plant provenance (i.e. native ecotypes vs. cultivars) was particularly important for the price. Seed mixtures containing only native ecotypes had considerably higher prices per ha (⩾+75%) than those including cultivars. In conclusion, increasing biodiversity in grasslands can be costly. These costs need to be considered when making recommendations to farmers and other stakeholders. Measures to reduce such costs for maintaining and/or increasing plant diversity could promote establishment of grasslands with higher plant diversity, facilitate the restoration of semi-natural grasslands, and contribute to solving the biodiversity crisis in agroecosystems.

1. Recent declines in biodiversity have given new urgency to questions about the relationship between land-use change, biodiversity and ecosystem processes. Despite the existence of a large body of research on the effects of land use on species richness, it is unclear whether the effects of land use on species richness are principally direct or indirect, mediated by concomitant changes in ecosystem processes. Therefore, we compared the direct effects of land use (fertilization, mowing and grazing) on species richness with indirect ones (mediated via grassland productivity) for grasslands in central Europe. 2. We measured the richness and above-ground biomass in 150 grassland plots in 3 regions of Germany (the so-called Biodiversity Exploratories). We used univariate and structural equation models to examine direct and indirect land-use effects. 3. The direct effects of mowing (-0.37, effect size) and grazing (0.04) intensity on species richness were stronger compared with the indirect effects of mowing (-0.04) and grazing (-0.01). However, the strong negative effect of fertilization (-0.23) on species richness was mainly indirect, mediated by increased productivity compared with the weak direct negative effect (-0.07). 4. Differences between regions in land-use effects showed five times weaker negative effects of mowing (-0.13) in the region with organic soils (Schorfheide-Chorin), strong overall negative effects of grazing (-0.29) for the region with organic soils opposed to a similar strong positive effect (0.30) in the Hainich-Dün region, whereas the Schwäbische Alb region displayed a five times weaker positive effect (0.06) only. Further, fertilization effects on species richness were positive (0.03) for the region with organic soils compared to up to 25 times stronger negative effects in the other two regions. 5. Synthesis. Our results clearly show the importance of studying both direct and indirect effects of land-use intensity. They demonstrate the indirect nature, via productivity, of the negative effect of fertilization intensity on plant species richness in the real-world context of management-induced gradients of intensity of fertilization, mowing and grazing. Finally, they highlight that careful consideration of regional environments is necessary before attempting to generalize land-use effects on species diversity.

1. Land-use intensification drives changes in microbial communities and the soil functions they regulate, but the mechanisms underlying these changes are poorly understood as land use can affect soil communities both directly (e.g. via changes in soil fertility) and indirectly (e.g. via changes in plant inputs). 2. The speed of microbial responses is also poorly understood. For instance, whether it is long-term legacies or short-term changes in land-use intensity that drive changes in microbial communities. 3. To address these topics, we measured multiple microbial functions, bacterial and fungal biomass and abiotic soil properties at two time intervals 3 years apart. This was performed in 150 grassland sites differing greatly in management intensity across three German regions. 4. Observed changes in microbial soil properties were related to both long-term means and short-term changes in: abiotic soil properties, land-use intensity, community abundance-weighted means of plant functional traits and plant biomass properties in regression and structural equation models. Plant traits, particularly leaf phosphorus, and soil pH were the best predictors of change in soil microbial function, as well as fungal and bacterial biomass, while land-use intensity showed weaker effects. 5. Indirect legacy effects, in which microbial change was explained by the effects of long-term land-use intensity on plant traits, were important, thus indicating a time lag between plant community and microbial change. Whenever the effects of short-term changes in land-use intensity were present, they acted directly on soil microorganisms. 6. Synthesis. The results provide new evidence that soil communities and their functioning respond to short-term changes in land-use intensity, but that both rapid and longer time-scale responses to changes in plant functional traits are at least of equal importance. This suggests that management which shapes plant communities may be an effective means of managing soil communities and the functions and services they provide.

Aims: We investigated urban grasslands to: (1) explore current patterns of plant species richness in high-maintenance vs low-maintenance grasslands, (2) investigate environmental drivers of plant species richness and composition, and (3) derive management recommendations and assess the potential for plant species introduction. Location: Cities of Cologne (50°56' N, 6°57' E) and Münster (51°57' N, 7°37' E), North Rhine-Westphalia, Germany. Methods: We performed plant inventories and measured soil and above-ground biomass characteristics in 100 urban grasslands in two cities differing in population size and environmental setting. The data set covered 35 high-maintenance grasslands, which are cut or mulched up to 14 times a year, and 65 lowmaintenance grasslands with one to two cuts per year or sheep grazing. We used ANCOVA and DCA to assess drivers of vegetation composition and species richness. The floristic potential and options to restore biodiversity were assessed taking into account maintenance intensity and key abiotic variables of the grasslands using thresholds derived from published literature and our own data. Results: High-maintenance urban grasslands harboured significantly lower plant species richness compared to low-maintenance grasslands. However, plant species richness of both grassland types turned out to be lower than that of comparable semi-natural agricultural grasslands. Floristic composition was primarily conditioned by maintenance intensity, but for plant species richness environmental factors such as soil pH, phosphorus availability and city were additionally important. Just eight of the 100 studied urban grasslands were found to be already valuable and species-rich, whereas the vast majority showed relatively low species richness but a high potential for species introduction. Conclusions: Apparently, most urban grasslands exhibited quite constrained plant species richness, suffering from high-maintenance intensity but probably also from dispersal and seed limitations. Nevertheless, as the majority of the studied grasslands showed favourable abiotic preconditions for higher plant species richness, restoration techniques using species introduction could be an easy and promising method to support grassland biodiversity in urban areas.

An important limitation of studies on the ecological value of urban habitats is the question of habitat connectivity. While connectivity allows spread and genetic exchange of species, isolation can lead to the extinction of populations through inbreeding and loss of genetic diversity. However, while practitioners already start improving urban habitat patches by means of restoration, issues of connectivity have not been taken in account although being crucial for a self-sustaining habitat network. In our study, we evaluate the connectivity of public urban grasslands such as lawns, grasslands in residential areas, playgrounds or parks in the city of Münster, Germany. Employing a graph theory approach, which uses a map (graph) of habitat patches to calculate functional connectivity measures using patch area size and patch connectivity, we studied four groups of different plant dispersal distances (2 m, 20 m, 44 m and 100 m). This approach can be recommended for planning issues due to the output of different indicator values and comparably low data demands. Our results show for the first time, that connectivity of urban grasslands is extremely low for species of short-distance dispersal but slightly increases for long-distance dispersal. Except two larger conglomerations of well-connected patches, urban grasslands were found to be mostly sparsely connected. Thus, future research should focus on options to improve connectivity and to prioritize patches for ecological improvement. The graph theory approach turned out to be a useful tool to analyse urban habitat connectivity but also to illustrate results obtained.

Nitrogen fertilisation is a common form of agricultural intensification, aimed at increasing biomass, which can affect plant species diversity and ecosystem functioning. Using a systematic review and meta-analysis of nitrogen fertilisation studies in European permanent grasslands, we asked: (i) what relationship form exists between nitrogen application rate and change in plant diversity, compared to zero fertilisation controls; and (ii) how grassland, management and study characteristics affect this relationship. Meta-analysis of 34 control-treatment effects from 14 studies conducted across nine European countries revealed a negative linear relationship between nitrogen fertilisation rate and change in plant species richness, equivalent to approximately 1.5 species/m2 lost for every 100 Kg ha−1 yr−1 of nitrogen added. Fertilisation induced reductions in plant species richness were greater when defoliation rates were lower. We found some evidence that grasslands with a higher baseline plant diversity lost more species when fertilised compared to more species poor grasslands, although uncertainty was high. Due to the diverse grassland types included in the analysis, the variability in fertilisation-driven changes in plant diversity was high. We identified several remaining limitations to our understanding, including uncertainty about non-linear effects, which could aid efforts to optimise the trade-off of plant diversity and increasing grassland yields.

While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m(2)) and central Germany (6.4 species per 16 m(2)) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.

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. Our analyses revealed land use, nutrient availability, plant functionality, and species richness driving plant community composition/structure in a complex manner. For deeper understanding of underlying mechanisms shaping community assembly and biodiversity conservation, we suggest the consideration of all these parameters.