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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.

Many studies have analysed the effect of browsing by large herbivores on tree species but far fewer studies have studied their effect on understorey shrubs and herbs. Moreover, while many studies have shown that forest features and management intensity strongly influence understorey vegetation, the influence of such variation on the effect of large-herbivore exclusion is not known. In this study, we analysed changes of species richness, Shannon diversity, evenness and cover of understorey herbs and shrubs after excluding large herbivores for seven years on 147 forest sites, differing in management intensity and forest features, in three regions of Germany (Schwäbische Alb, Hainich-Dün, Schorfheide-Chorin). Further, we studied how the effect of large-herbivore exclusion on understorey vegetation was influenced by forest management intensity and several forest features. As expected, exclusion of large herbivores resulted in highly variable results. Nevertheless, we found that large-herbivore exclusion significantly increased cover and Shannon diversity of shrub communities, while it did not affect herb communities. Forest management intensity did not influence the effect of large-herbivore exclusion while some forest features, most often relative conifer cover, did. In forests with high relative conifer cover, large-herbivore exclusion decreased species richness and cover of herbs and increased Shannon diversity of herbs and shrubs, while in forests with low relative conifer cover large-herbivore exclusion increased species richness and cover of herbs, and decreased Shannon diversity of herbs and shrubs. We suggest that browsing by large herbivores should be included when studying understorey shrub communities, however when studying understorey herb communities the effects of browsing are less general and depend on forest features.

Aim Despite increasing interest in β‐diversity, that is the spatial and temporal turnover of species, the mechanisms underlying species turnover at different spatial scales are not fully understood, although they likely differ among different functional groups. We investigated the relative importance of dispersal limitations and the environmental filtering caused by vegetation for local, multi‐taxa forest communities differing in their dispersal ability, trophic position and body size. Location Temperate forests in five regions across Germany. Methods In the inter‐region analysis, the independent and shared effects of the regional spatial structure (regional species pool), landscape spatial structure (dispersal limitation) and environmental factors on species turnover were quantified with a 1‐ha grain across 11 functional groups in up to 495 plots by variation partitioning. In the intra‐region analysis, the relative importance of three environmental factors related to vegetation (herb and tree layer composition and forest physiognomy) and spatial structure for species turnover was determined. Results In the inter‐region analysis, over half of the explained variation in community composition (23% of the total explained 35%) was explained by the shared effects of several factors, indicative of spatially structured environmental filtering. Among the independent effects, environmental factors were the strongest on average over 11 groups, but the importance of landscape spatial structure increased for less dispersive functional groups. In the intra‐region analysis, the independent effect of plant species composition had a stronger influence on species turnover than forest physiognomy, but the relative importance of the latter increased with increasing trophic position and body size. Main conclusions Our study revealed that the mechanisms structuring assemblage composition are associated with the traits of functional groups. Hence, conservation frameworks targeting biodiversity of multiple groups should cover both environmental and biogeographical gradients. Within regions, forest management can enhance β‐diversity particularly by diversifying tree species composition and forest physiognomy.

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.

Background Temperate forest understorey vegetation poses an excellent study system to investigate whether increases in resource availability lead to an increase in plant species richness. Most sunlight is absorbed by the species-poor tree canopy, making the much more species-rich understorey species inhabit a severely resource-limited habitat. Additionally, the heterogeneity of light availability, resulting from management-moderated tree composition and age structure, may contribute to species coexistence. One would therefore expect that the diversity in the herb layer correlates positively with either the overall light availability, or the light heterogeneity, depending on whether resource availability or heterogeneity are more important drivers of diversity. To test this idea, we assessed variability of light conditions in 75 forest plots across three ecoregions with four different methods. Results We correlated these data with vegetation relevés and found light availability to be strongly positively correlated with understorey plant species richness, as well as with understorey cover. Light variability (assessed with two approaches) within plots was positively correlated with transmittance, but did not improve the relationship further, suggesting that the main driver of species richness in this system is the overall resource availability. Two of the three beech-dominated regions exhibited near-identical effects of light transmittance, while the third, featuring pine alongside beech and thus with the longest gradient of transmittance and lowest species richness, displayed a weaker light response. Conclusions While site conditions are certainly responsible for the trees selected by foresters, for the resulting forest structure, and for the differences in plant species pools, our results suggest that light transmittance is a strong mediating factor of understorey plant species richness.

Aim Intensification of land use strongly impacts plant communities by causing shifts in taxonomic and functional composition. Mechanisms of land use‐induced biodiversity losses have been described for temperate grasslands, but a quantitative assessment of species‐specific occurrence optima and maximum tolerance (niche breadth) to land‐use intensity (LUI) in Central European grasslands is still lacking. Location Temperate, managed permanent grasslands in three regions of Germany. Methods We combined extensive field work with a null model–randomization approach, defined a “habitat niche” for each plant species based on occurrence and abundance across 150 grassland sites differing in LUI (i.e., amount of fertilizer, mowing/grazing intensity and a compound index of these), and assessed their realized niche breadth (tolerance). Underlying mechanisms driving species’ responses to LUI were assessed by relating plant functional traits, Ellenberg indicator values (EIV), Grime's ecological strategies (CSR) and Briemle utilization numbers. Results Out of 151 plant species, 34% responded negatively, whereas 10% responded positively to high LUI. This pattern was mainly driven by species’ response to fertilization and mowing frequency; grazing intensity response was less pronounced. Positively reacting species, displaying broader niches, were associated with competition‐related functional traits, high EIV for nutrient supply and moisture and high mowing tolerance under spatiotemporally variable conditions. Negatively responding species, displaying relatively narrow niches confined to spatiotemporally homogeneous low LUI sites, were associated with a nutrient‐retentive strategy, under nutrient‐poor, base‐rich soil conditions. Conclusion Our analyses of individual species’ reactions clearly demonstrate that species responding negatively to high LUI display little tolerance towards intensive fertilization and mowing, leading to plant diversity loss; whereas grazing partly thwarts these effects by creating new habitat niches and promoting ruderal species. Our approach can be applied to other habitat types and biogeographical regions in order to quantify local specific response or tolerance, adding to existing knowledge about local vegetation dynamics. Employing a combined null model–randomization approach based on species’ occurrence and abundance in temperate grasslands, we calculated plant species‐specific agricultural habitat niches and niche breadths and characterized response‐driving mechanisms by using plant functional and ecological traits. Our approach may be applied to any other habitat type for explaining and predicting community assembly and species coexistence in response to current land‐use practices.

Land‐use intensification is a major driver of local species extinction and homogenization. Temperate grasslands, managed at low intensities over centuries harbored a high species diversity, which is increasingly threatened by the management intensification over the last decades. This includes key taxa like ants. However, the underlying mechanisms leading to a decrease in ant abundance and species richness as well as changes in functional community composition are not well understood. We sampled ants on 110 grassland plots in three regions in Germany. The sampled grasslands are used as meadows or pastures, being mown, grazed or fertilized at different intensities. We analyzed the effect of the different aspects of land use on ant species richness, functional trait spaces, and community composition by using a multimodel inference approach and structural equation models. Overall, we found 31 ant species belonging to 8 genera, mostly open habitat specialists. Ant species richness, functional trait space of communities, and abundance of nests decreased with increasing land‐use intensity. The land‐use practice most harmful to ants was mowing, followed by heavy grazing by cattle. Fertilization did not strongly affect ant species richness. Grazing by sheep increased the ant species richness. The effect of mowing differed between species and was strongly negative for Formica species while Myrmica and common Lasius species were less affected. Rare species occurred mainly in plots managed at low intensity. Our results show that mowing less often or later in the season would retain a higher ant species richness—similarly to most other grassland taxa. The transformation from (sheep) pastures to intensively managed meadows and especially mowing directly affects ants via the destruction of nests and indirectly via loss of grassland heterogeneity (reduced plant species richness) and increased soil moisture by shading of fast‐growing plant species. In our study, we analyzed the effect of land‐use intensity, mowing, fertilization, and grazing on temperate ant communities. Our results show that ant species respond sensitively to land‐use intensification which reduces species richness, abundance of nests, and functional trait spaces. The land‐use practice most harmful to ants was mowing, followed by heavy grazing by cattle.

1. Drivers of ecosystem stability have been a major topic in ecology for decades. Most studies have focused on the influence of species richness on ecosystem stability and found positive diversity-stability relationships. However, land use and abiotic factors shape species richness and functional composition of plant communities and may override species richness-stability relations in managed grasslands. 2. We analysed the relative importance of land-use intensity (LUI), resident plant species richness and functional composition for recovery of plant communities (plant species richness, plant cover, above- and below-ground biomass) and release of soil nutrients after a severe mechanical disturbance. Experimental sward disturbance was applied to 73 grassland sites along a LUI gradient in three German regions. We considered relative (ln(disturbance/control)) and absolute (disturbance - control) treatment effects. Using structural equation modelling, we disentangled direct effects of LUI and resident species richness on recovery and indirect effects via changes in functional richness. 3. Community-weighted-mean traits rarely mattered for recovery or nutrient release, while functional richness especially increased relative recovery of plant communities but also relative release of NO₃-N and NH₄-N. These effects were enhanced by increasing resident plant species richness and decreasing LUI. Next to these indirect influences of LUI and resident plant species richness via functional community composition, grasslands of high compared with grasslands of low resident plant species richness generally showed decreased recovery of plant communities. In grasslands of high LUI, absolute recovery of some aspects of plant communities was decreased. We did not find consistent differences between the relative importance of the different drivers of recovery after the first and the second season. Overall, resident species richness seemed most important for relative recovery and less important for absolute recovery, where direct effects of LUI were more common. 4. Synthesis. The stability of ecosystems in managed grasslands depends on more than species richness. Thus, drivers that directly affect species richness and functional community composition have to be considered when studying the stability of real-world ecosystems. More specifically, in managed grasslands high resident species richness but also high land-use intensity (LUI) decreased the stability of ecosystem functions, which was partially buffered by increases in functional richness.

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. Relationships between biodiversity and phosphorus cycling and the underlying processes are complex. Here the authors analyse a biodiversity manipulation experiment and an agricultural management gradient to show how plant and mycorrhizal fungal diversity promote phosphorus exploitation.

A large body of research shows that biodiversity loss can reduce ecosystem functioning, thus providing support for the conservation of biological diversity. Much of the evidence for this relationship is drawn from biodiversity-ecosystem functioning experiments (hereafter: biodiversity experiments), in which biodiversity loss is simulated by randomly assembling communities of varying species diversity, and ecosystem functions are measured. This random assembly has led some ecologists to question the relevance of biodiversity experiments to real-world ecosystems, where community assembly may often be non-random and influenced by external drivers, such as climate or land-use intensification. Despite these repeated criticisms, there has been no comprehensive, quantitative assessment of how experimental and real-world plant communities really differ, and whether these differences invalidate the experimental results. Here, we compare data from two of the largest and longest-running grassland biodiversity experiments globally (Jena Experiment, Germany; BioDIV, USA) to related real-world grassland plant communities in terms of their taxonomic, functional, and phylogenetic diversity and functional-trait composition. We found that plant communities of biodiversity experiments have greater variance in these compositional features than their real-world counterparts, covering almost all of the variation of the real-world communities (82-96%) while also containing community types that are not currently observed in the real world. We then re-analysed a subset of experimental data that included only ecologically-realistic communities, i.e. those comparable to real-world communities. For ten out of twelve biodiversity-ecosystem functioning relationships, biodiversity effects did not differ significantly between the full dataset of biodiversity experiments and the ecologically-realistic subset of experimental communities. This demonstrates that the results of biodiversity experiments are largely insensitive to the inclusion/exclusion of unrealistic communities. By bridging the gap between experimental and real-world studies, these results demonstrate the validity of inferences from biodiversity experiments, a key step in translating their results into specific recommendations for real-world biodiversity management.