Biomass removal promotes plant diversity after short-term de-intensification of managed grasslands.

Land-use intensification is one of the main drivers threatening biodiversity in managed grasslands. Despite multiple studies investigating the effect of different land-use components in driving changes in plant biodiversity, their effects are usually studied in isolation. Here, we establish a full factorial design crossing fertilization with a combined treatment of biomass removal, on 16 managed grasslands spanning a gradient in land-use intensity, across three regions in Germany. Specifically, we investigate the interactive effects of different land-use components on plant composition and diversity using structural equation modelling. We hypothesize that fertilization and biomass removal alter plant biodiversity, directly and indirectly, mediated through changes in light availability. We found that, direct and indirect effects of biomass removal on plant biodiversity were larger than effects of fertilization, yet significantly differed between season. Furthermore, we found that indirect effects of biomass removal on plant biodiversity were mediated through changes in light availability, but also by changes in soil moisture. Our analysis thus supports previous findings, that soil moisture may operate as an alternative indirect mechanism by which biomass removal may affect plant biodiversity. Most importantly, our findings highlight that in the short-term biomass removal can partly compensate the negative effects of fertilization on plant biodiversity in managed grasslands. By studying the interactive nature of different land-use drivers we advance our understanding of the complex mechanisms controlling plant biodiversity in managed grasslands, which ultimately may help to maintain higher levels of biodiversity in grassland ecosystems

Plant diversity effects on forage quality, yield and revenues of semi-natural grasslands

In agricultural settings, plant diversity is often associated with low biomass yield and forage quality, while biodiversity experiments typically find the opposite. We address this controversy by assessing, over 1 year, plant diversity effects on biomass yield, forage quality (i.e. nutritive values), quality-adjusted yield (biomass yield × forage quality), and revenues across different management intensities (extensive to intensive) on subplots of a large-scale grassland biodiversity experiment. Plant diversity substantially increased quality-adjusted yield and revenues. These findings hold for a wide range of management intensities, i.e., fertilization levels and cutting frequencies, in semi-natural grasslands. Plant diversity was an important production factor independent of management intensity, as it enhanced quality-adjusted yield and revenues similarly to increasing fertilization and cutting frequency. Consequently, maintaining and reestablishing plant diversity could be a way to sustainably manage temperate grasslands

Effects of Plant Diversity, Functional Group Composition, and Fertilization on Soil Microbial Properties in Experimental Grassland

Background: Loss of biodiversity and increased nutrient inputs are two of the most crucial anthropogenic factors driving ecosystem change. Although both received considerable attention in previous studies, information on their interactive effects on ecosystem functioning is scarce. In particular, little is known on how soil biota and their functions are affected by combined changes in plant diversity and fertilization. Methodology/principal findings: We investigated the effects of plant diversity, functional community composition, and fertilization on the biomass and respiration of soil microbial communities in a long-term biodiversity experiment in semi-natural grassland (Jena Experiment). Plant species richness enhanced microbial basal respiration and microbial biomass, but did not significantly affect microbial specific respiration. In contrast, the presence of legumes and fertilization significantly decreased microbial specific respiration, without altering microbial biomass. The effect of legumes was superimposed by fertilization as indicated by a significant interaction between the presence of legumes and fertilization. Further, changes in microbial stoichiometry (C-to-N ratio) and specific respiration suggest the presence of legumes to reduce N limitation of soil microorganisms and to modify microbial C use efficiency. Conclusions/significance: Our study highlights the role of plant species and functional group diversity as well as interactions between plant community composition and fertilizer application for soil microbial functions. Our results suggest soil microbial stoichiometry to be a powerful indicator of microbial functioning under N limited conditions. Although our results support the notion that plant diversity and fertilizer application independently affect microbial functioning, legume effects on microbial N limitation were superimposed by fertilization, indicating significant interactions between the functional composition of plant communities and nutrient inputs for soil processes

Plant diversity influenced gross nitrogen mineralization, microbial ammonium consumption and gross inorganic N immobilization in a grassland experiment

Gross rates of nitrogen (N) turnover inform about the total N release and consumption. We investigated how plant diversity affects gross N mineralization, microbial ammonium (NH4+) consumption and gross inorganic N immobilization in grasslands via isotopic pool dilution. The field experiment included 74 plots with 1–16 plant species and 1–4 plant functional groups (legumes, grasses, tall herbs, small herbs). We determined soil pH, shoot height, root, shoot and microbial biomass, and C and N concentrations in soil, microbial biomass, roots and shoots. Structural equation modeling (SEM) showed that increasing plant species richness significantly decreased gross N mineralization and microbial NH4+ consumption rates via increased root C:N ratios. Root C:N ratios increased because of the replacement of legumes (low C:N ratios) by small herbs (high C:N ratios) and an increasing shoot height, which was positively related with root C:N ratios, with increasing species richness. However, in our SEM remained an unexplained direct negative path from species richness to both N turnover rates. The presence of legumes increased gross N mineralization, microbial NH4+ consumption and gross inorganic N immobilization rates likely because of improved N supply by N2 fixation. The positive effect of small herbs on microbial NH4+ consumption and gross inorganic N immobilization could be attributed to their increased rhizodeposition, stimulating microbial growth. Our results demonstrate that increasing root C:N ratios with increasing species richness slow down the N cycle but also that there must be additional, still unidentified processes behind the species richness effect potentially including changed microbial community composition

Drought-exposure history increases complementarity between plant species in response to a subsequent drought

Growing threats from extreme climatic events and biodiversity loss have raised concerns about their interactive consequences for ecosystem functioning. Evidence suggests biodiversity can buffer ecosystem functioning during such climatic events. However, whether exposure to extreme climatic events will strengthen the biodiversity-dependent buffering effects for future generations remains elusive. We assess such transgenerational effects by exposing experimental grassland communities to eight recurrent summer droughts versus ambient conditions in the field. Seed offspring of 12 species are then subjected to a subsequent drought event in the glasshouse, grown individually, in monocultures or in 2-species mixtures. Comparing productivity between mixtures and monocultures, drought-selected plants show greater between-species complementarity than ambient-selected plants when recovering from the subsequent drought, causing stronger biodiversity effects on productivity and better recovery of drought-selected mixtures after the drought. These findings suggest exposure to recurrent climatic events can improve ecosystem responses to future events through transgenerational reinforcement of species complementarity

Relationships between species richness and biomass production are context dependent in grasslands differing in land-use and seed addition.

Despite evidence from grasslands experiments suggesting that plant species loss reduces biomass production, the strength of biodiversity-ecosystem functioning relationships in managed grasslands is still debated. High land-use intensity and reduced species pools are often suggested to make relationships between biodiversity and productivity less positive or even negative, but concrete evidence is still scarce. We investigated biodiversity-productivity relationships over two years in 150 managed grasslands in Germany. Specifically, we distinguished between relationships of biodiversity and biomass production in managed grasslands (1) varying in land-use intensity (e.g. of mowing, grazing and/or fertilization), (2) where land-use intensity is experimentally reduced, and (3) where additionally to land-use reductions, species pools are enlarged by seed addition. Among grasslands varying in land-use intensity, we found negative biodiversity-productivity relationships. Land-use reduction weakened these relationships, towards neutral, and sometimes, even positive relationships. Seed addition reduced species pool limitations, but this did not strengthen biodiversity-productivity relationships. Our findings indicate that land-use intensity is an important factor explaining the predominantly negative biodiversity-productivity relationships in managed grasslands. While we did not find that species pool limitations weakened biodiversity-productivity relationships, our results are based on a two-year-old experiment, possibly such effects are only visible in the long-term. Ultimately, advancing insights on biodiversity-ecosystem functioning relationships helps us to understand under which conditions agricultural production may benefit from promoting biodiversity

Trait means, trait plasticity and trait differences to other species jointly explain species performances in grasslands of varying diversity

Functional traits may help to explain the great variety of species performances in plant communities, but it is not clear whether the magnitude of trait values of a focal species or trait differences to co‐occurring species are key for trait‐based predictions. In addition, trait expression within species is often plastic, but this variation has been widely neglected in trait‐based analyses. We studied functional traits and plant biomass of 59 species in 66 experimental grassland mixtures of varying species richness (Jena Experiment). We related mean species performances (species biomass and relative yield RY) and their plasticities along the diversity gradient to trait‐based pedictors involving mean species traits (Tmean), trait plasticities along the diversity gradient (Tslope), extents of trait variation across communities (TCV; coefficient of variation) and hierarchical differences (Tdiff) and trait distances (absolute values of trait differences Tdist) between focal and co‐occurring species. Tmean (30–55%) and Tdiff (30–33%) explained most variation in mean species performances and their plasticities, but Tslope (20–25%) was also important in explaining mean species performances. The mean species traits and the trait differences between focal species and neighbors with the greatest explanatory power were related to plant size and stature (shoot length, mass:height ratios) and leaf photosynthetic capacity (specific leaf area, stable carbon isotopes and leaf nitrogen concentration). The contribution of trait plasticities in explaining species performances varied in direction (positive or negative) and involved traits related to photosynthetic capacity, nitrogen acquisition (nitrogen concentrations and stable isotopes) as well as structural stability (shoot carbon concentrations). Our results suggest that incorporating plasticity in trait expression as well as trait differences to co‐occurring species is critical for extending trait‐based analyses to understand the assembly of plant communities and the contribution of individual species in structuring plant communities