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Root-associated fungi could play a role in determining both the positive relationship between plant diversity and productivity in experimental grasslands, and its strengthening over time. This hypothesis assumes that specialized pathogenic and mutualistic fungal communities gradually assemble over time, enhancing plant growth more in species-rich than in species-poor plots. To test this hypothesis, we used high-throughput amplicon sequencing to characterize root-associated fungal communities in experimental grasslands of 1 and 15 years of age with varying levels of plant species richness. Specifically, we tested whether the relationship between fungal communities and plant richness and productivity becomes stronger with the age of the experimental plots. Our results showed that fungal diversity increased with plant diversity, but this relationship weakened rather than strengthened over the two time points. Contrastingly, fungal community composition showed increasing associations with plant diversity over time, suggesting a gradual build-up of specific fungal assemblages. Analyses of different fungal guilds showed that these changes were particularly marked in pathogenic fungi, whose shifts in relative abundance are consistent with the pathogen dilution hypothesis in diverse plant communities. Our results suggest that root-associated fungal pathogens play more specific roles in determining the diversity–productivity relationship than other root-associated plant symbionts.

Eutrophication usually impacts biodiversity, species composition, and functioning of grassland communities. Whether such effects propagate to influence the stability of these community aspects is unknown. Using standardized experiments across 55 global grasslands, we quantified the effects of nutrient addition on five stability facets (i.e., temporal invariability and resistance during and recovery after dry and wet growing seasons) for three community aspects (i.e., aboveground biomass, community composition, and species richness). Nutrient addition reduced the temporal invariability and resistance of species richness and community composition, but not biomass, during dry and wet growing seasons. Temporal invariability and resistance during, but not recovery after, dry and wet growing seasons were strongly positively correlated in both ambient and eutrophic conditions. This indicates that maintaining and restoring the stability of plant communities requires increasing resistance rather than recovery. Harnessing the complexity of ecological stability provides new insights for grassland ecosystem sustainability in a changing world.

Ecosystem management aims at providing many ecosystem services simultaneously. Such ecosystem multifunctionality can be limited by trade-offs and increased by synergies among the underlying ecosystem functions (EF), which need to be understood to develop targeted management. Previous studies found differences in the correlation between EFs. We hypothesised that correlations between EFs are variable even under the controlled conditions of a field experiment and that seasonal and annual variation, plant species richness, and plot identity (identity effects of plant communities such as the presence and absence of functional groups and species) are drivers of these correlations. We used data on 31 EFs related to plants, consumers, and physical soil properties that were measured over 5 to 19 years, up to three times per year, in a temperate grassland experiment with 80 different plots, constituting six sown plant species richness levels (1, 2, 4, 8, 16, 60 species). We found that correlations between pairs of EFs were variable, and correlations between two particular EFs could range from weak to strong correlations or from negative to positive correlations among the repeated measurements. To determine the drivers of pairwise EF correlations, the covariance between EFs was partitioned into contributions from plant species richness, plot identity, and time (including years and seasons). We found that most of the covariance for synergies was explained by species richness (26.5%), whereas for trade-offs, most covariance was explained by plot identity (29.5%). Additionally, some EF pairs were more affected by differences among years and seasons and therefore showed a higher temporal variation. Therefore, correlations between two EFs from single measurements are insufficient to draw conclusions on trade-offs and synergies. Consequently, pairs of EFs need to be measured repeatedly under different conditions to describe their relationships with more certainty and be able to derive recommendations for the management of grasslands.

Extreme weather events are occurring more frequently, and research has shown that plant diversity can help mitigate impacts of climate change by increasing plant productivity and ecosystem stability1,2. Although soil temperature and its stability are key determinants of essential ecosystem processes related to water and nutrient uptake3as well as soil respiration and microbial activity4, no study has yet investigated whether plant diversity can buffer soil temperature fluctuations. Using 18 years of a continuous dataset with a resolution of 1 minute (∼795,312,000 individual measurements) from a large-scale grassland biodiversity experiment, we show that plant diversity buffers soil temperature throughout the year. Plant diversity helped to prevent soil heating in hot weather, and cooling in cold weather. Moreover, this effect of plant diversity increased over the 18-year observation period with the aging of experimental communities and was even stronger under extreme conditions, i.e., on hot days or in dry years. Using structural equation modelling, we found that plant diversity stabilized soil temperature by increasing soil organic carbon concentrations and, to a lesser extent, by increasing the plant leaf area index. We suggest that the diversity-induced stabilization of soil temperature may help to mitigate the negative effects of extreme climatic events such as soil carbon release, thus slow global warming.

Plant monocultures growing for extended periods face severe losses of productivity. This phenomenon, known as ‘yield decline’, is often caused by the accumulation of above- and belowground plant antagonists. The effectiveness of plant defences against antagonists might help explaining differences in yield decline among species. Using a trait-based approach, we studied the role of 20 physical and chemical defence traits of leaves and fine roots on yield decline of 18-year old monocultures of 27 grassland species. We hypothesized that yield decline is lower for species with high defences, that root defences are better predictors of yield decline than leaf defences, and that in roots, physical defences better predict yield decline than chemical defences, while the reverse is true for leaves. We additionally hypothesized that species increasing the expression of defence traits after long-term monoculture growth would suffer less yield decline. We summarized leaf and fine root defence traits using principal component analysis and analysed the relationship between defence traits mean as a measure of defence strenght and defence traits temporal changes of the most informative components and monoculture yield decline. The only significant predictors of yield decline were the mean and temporal changes of the component related to specific root length and root diameter (e.g. the so called collaboration gradient of the root economics space). The principal component analysis of the remaining traits showed strong trade-offs between defences suggesting that different plant species deploy a variety of strategies to defend themselves. This diversity of strategies could preclude the detection of a generalized correlation between the strength and temporal changes of defence gradients and yield decline. Our results show that yield decline is strongly linked to belowground processes particularly to root traits. Further studies are needed to understand the mechanism driving the effect of the collaboration gradient on yield decline.

There is no doubt about the insect decline currently taking place in ecosystems with large anthropogenic impacts. Thus, there is a need for practices that avoid insect decline and or help to recover insect communities that have already suffered. Plant diversity has been shown to be positively related to insect abundance and diversity and to ecosystem functions provided by insects. However, it remains open if increased plant diversity can help to recover decreased populations. Here, we tested over one decade the effects of plant diversity on the carabid community in a large grassland biodiversity experiment and how plant diversity fostered the establishment of a natural grassland community after conversion of an arable field. There was a dramatic decline in carabid abundance from 2003, the first year after establishing the diversity experiment, to 2005. However, subsequently, the abundance increased constantly. One year after the land use change most individuals and species were those commonly found in agricultural fields. In subsequent years the community was dominated by grassland species. While plant diversity did not affect the abundance and richness of the carabid community, the turnover to a more native grassland community was accelerated by plant diversity in the first years after the land use change. In contrast, in later years plant diversity stabilized the community assemblage. Our study shows that high plant diversity can contribute to a faster transition of insect populations towards naturally occurring community assemblages and at later stages to more stabilized assemblages.

Seed removal can result in either seed predation or dispersal and is therefore an indicator of important ecosystem functions. To better understand how these variable processes are affected by environmental changes, rapid and reliable assessments with high temporal and spatial replication are needed. We address this need by testing the application of a rapid ecosystem function assessment (REFA) method to investigate seed removal in habitats with differing land use intensity. We recorded seed removal hourly over eight hours at 301 sites in five habitat types in three urban regions in Germany. We calculated seed removal rates per sampling period, traditionally used in seed removal studies, as well as instantaneous seed removal probabilities based on hourly sampling. Across regions, seed removal probabilities and rates were lowest in arable fields, a habitat type with high land-use intensity. Except in urban sealed areas, temperature had a negative effect on seed removal. Additional Cox hazard regressions suggest invertebrates as the main seed removing animals in arable fields, whereas vertebrates were likely removers in other habitat types. We confirm that seed removal is strongly negatively affected by human disturbance, indicating that the tested method is appropriate in different settings. We were able to recognize patterns in highly variable data and the method also has the advantages of low cost, high replication and high temporal resolution. However, there is a trade-off between the high temporal-resolution of instantaneous seed removal probabilities and the sampling effort, but adjustments in the standardized setup can be made depending on the study. To further utilize the extensive data collection in the REFA method, we propose to combine instantaneous seed removal probabilities, seed removal rates and Cox hazard regressions of seed removal to provide complementary information on the extent and temporal patterns of seed removal and indications about potential seed removing guilds. Keywords: Seed predation, seed dispersal, seed removal, granivore, REFA

SummaryEutrophication impacts grassland diversity such as species richness, functional trait diversity and composition, but whether and how these changes affect ecosystem functional stability under increasing climate extremes is unknown. We quantify the direct and diversity-mediated effects of nutrient addition on functional stability of productivity as measured by resistance during and recovery after dry and wet growing seasons, and temporal invariability across 19 grasslands spanning four continents. We use leaf traits measured in the field to construct slow-fast leaf economic spectrum and calculate functional trait composition and diversity of the slow-fast trait. Nutrient addition increased resistance during dry growing seasons mainly through its direct effects. It also increased resistance during wet growing seasons directly, but these positive effects were weakened by indirect negative effects through reducing slow-fast functional diversity. Nutrient addition had weak effects on recovery after dry growing seasons, but it decreased recovery after wet growing seasons mainly through its direct effects. Nutrient addition increased temporal invariability through resistance, but decreased it through recovery and reducing species richness, leading to overall weak net effects. Thus, processes and influencing pathways independent of plant diversity should be carefully taken into account in understanding ecosystem stability in response to increasing global changes.

Numerous studies have demonstrated that biodiversity drives ecosystem functioning, yet there is a lack of knowledge about how biodiversity loss alters ecosystems functioning and stability in the long-term. We report on temporal changes in the species richness–productivity, –stability, –species asynchrony, and –complementarity relationships over 17 years in a grassland biodiversity experiment. Richness–productivity and richness–complementarity relationships strengthened because productivity declined more rapidly in less than in more diverse communities. Using five-year rolling windows, we found that richness–stability relationships became stronger over time because more diverse communities maintained greater productivity. In later years asynchrony increased temporal complementarity among species, thus providing an insurance effect of species richness on stability. These results show how mechanisms stabilizing ecosystem functioning can change with community age. They demonstrate the importance of preserving diversity in natural systems as the role of diversity for ecosystem stability may take decades to realize the full potential.

A large body of research shows that biodiversity loss can reduce ecosystem functioning. However, much of the evidence for this relationship is drawn from biodiversity-ecosystem functioning 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 or disassembly may be non-random and influenced by external drivers, such as climate, soil conditions or land use. Here, we compare data from real-world grassland plant communities with data from two of the largest and longest-running grassland biodiversity experiments (the Jena Experiment in Germany and BioDIV in the United States) in terms of their taxonomic, functional and phylogenetic diversity and functional-trait composition. We found that plant communities of biodiversity experiments cover almost all of the multivariate variation of the real-world communities, while also containing community types that are not currently observed in the real world. Moreover, they have greater variance in their compositional features than their real-world counterparts. We then re-analysed a subset of experimental data that included only ecologically realistic communities (that is, those comparable to real-world communities). For 10 out of 12 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. Although we do not provide direct evidence for strong or consistent biodiversity-ecosystem functioning relationships in real-world communities, our results demonstrate that the results of biodiversity experiments are largely insensitive to the exclusion of unrealistic communities and that the conclusions drawn from biodiversity experiments are generally robust. By comparing data from real-world grassland communities with data from two of the longest-running grassland biodiversity-ecosystem functioning experiments, the authors show that conclusions derived from experimental systems are robust to the removal of unrealistic experimental communities., Nature Ecology & Evolution, 4 (11), ISSN:2397-334X

Global change drivers such as anthropogenic nutrient inputs simultaneously alter biodiversity, species composition, and ecosystem functions such as above ground biomass. These changes are interconnected by complex feedbacks among extinction, invasion, and shifting relative abundance. Here, we use a novel temporal application of the Price equation to separate species richness and biomass change through time and quantify the functional contributions of species that are lost, gained, and persist under ambient and experimental nutrient addition in 59 global grasslands. Under ambient conditions, compositional and biomass turnover was high, but species losses (i.e., local extinctions) were balanced by gains (i.e. colonization). Under fertilization, there was biomass loss associated with species loss. Few species were gained in fertilized conditions over time but those that were, and species that persisted, contributed to net biomass gains, outweighing biomass loss. These components of community change are associated with distinct effects on measures of ecosystem functioning.

Knowledge from agriculture and ecological field studies suggests that plant monocultures lose productivity over time, but the drivers underlying the long‐term performance of monocultures of grassland species are not completely understood. We examined the performance of 60 grassland species growing in monoculture for 12 years in a biodiversity experiment (Jena Experiment) and studied three groups of biotic drivers potentially affecting plant performance in monocultures over time: 1) soil biota (nematode communities, arbuscular mycorrhizal fungi), 2) leaf traits related to leaf economics spectrum, and 3) plant life‐cycle characteristics related to buffered population growth (viable seeds in topsoil, seedling density, seed survival). Monocultures of 15 out of 60 species increased productivity, while monocultures of the remaining 45 species showed slight to strong losses of productivity over time, resulting in zero biomass in 15 species. All three biotic drivers were related to the varying long‐term performance of monocultures. Their combined influence on monoculture performance could be interpreted as a tradeoff between ‘fast’ versus ‘slow’ life strategies. ‘Fast’ species showed rapid resource use and little buffering of population growth through a viable seed bank, which led to high biomass production in young monocultures but a consecutive loss of biomass production over time. ‘Slow’ species were characterized by positive nematode effects (high abundance of predatory nematodes), conservative use of resources, and a viable seed bank with high recruitment success resulting in gradually increasing productivity over time. In summary, our study highlights the importance of studying long‐term field monocultures to investigate the complex role of different biotic drivers responsible for productivity changes over time. These insights provide an essential baseline for estimating biodiversity effects on productivity as well as to understand and predict long‐term performance of plant populations.

Species‐rich plant communities can induce unique soil biotic legacy effects through changing the abundance and composition of soil biota. These soil legacy effects can cause feedbacks to influence plant performance. In addition, soil biota can induce (defensive) secondary metabolites in shoots and roots and thus affect plant–herbivore interactions. We hypothesize that plant diversity‐driven soil biotic legacy effects elicit changes in the shoot and root metabolome. We tested this hypothesis by establishing an experiment with four plant species. We grew plants in a sterile substrate inoculated with soil conditioned by different plant species communities: (a) monocultures of either of the four species, (b) the four species in a mixture, (c) an eight species mixture including all four species or (d) a sterile inoculum. After at least 8 weeks in the field, we estimated shoot herbivory. At the same time, we took root and shoot samples for metabolomics analyses by liquid chromatography quadrupole time‐of‐flight mass spectrometry. We found that shoot and root metabolomes of all plants grown in sterile soil differed significantly from those grown in living soil. The plant metabolomes in living soils differed by species and tissue. Across all species, shoots displayed a greater richness of secondary metabolites than roots. The richness of secondary metabolites differed by species and among living soils. The conditioning species richness significantly affected the Shannon diversity of secondary metabolites in Centaurea jacea. Shoot herbivory positively correlated with the richness and Shannon diversity of secondary metabolites in Leucanthemum vulgare. We detected multiple metabolites that together explained up to 88% of the variation in herbivory in the shoots of C. jacea and Plantago lanceolata. Synthesis.

1. Plant-soil feedback (PSF) has gained attention as a mechanism promoting plant growth and coexistence. However, because most PSF research has measured monoculture growth in greenhouse conditions, field-based PSF experiments remain an important frontier for PSF research. 2. Using a four-year, factorial field experiment in Jena, Germany, we measured the growth of nine grassland species on soils conditioned by each of the target species (i.e., PSF). Plant community models were parameterized with or without these PSF effects, and model predictions were compared to plant biomass production in new and existing diversity-productivity experiments. 3. Plants created soils that changed subsequent plant biomass by 36%. However, because they were both positive and negative, the net PSF effect was 14% less growth on ‘home’ than ‘away’ soils. At the species level, seven of nine species realized non-neutral PSFs, but the two dominant species grew only 2% less on home than away soils. At the species*soil type level, 31 of 72 PSFs differed from zero. 4. In current and pre-existing diversity-productivity experiments, nine-species plant communities produced 37 to 29% more biomass than monocultures due primarily to selection effects. Null and PSF models predicted 29 to 28% more biomass for polycultures than monocultures, again due primarily to selection effects. 5. Synthesis: In field conditions, PSFs were large enough to be expected to cause roughly 14% overyielding due to complementarity, however, in plant communities overyielding was caused by selections effects, not complementarity effects. Further, large positive and large negative PSFs were associated with subdominant species, suggesting there may be selective pressure for plants to create neutral PSF. Broadly, results highlighted the importance of testing PSF effects in communities because there are several ways in which PSFs may be more or less important to plant growth in communities than suggested from simple PSF values.

Understanding factors that maintain ecosystem stability is critical in the face of environmental change. Experiments simulating species loss from grassland have shown that losing biodiversity decreases ecosystem stability. However, as the originally sown experimental communities with reduced biodiversity develop, plant evolutionary processes or the assembly of interacting soil organisms may allow ecosystems to increase stability over time. We explored such effects in a long‐term grassland biodiversity experiment with plant communities with either a history of co‐occurrence (selected communities) or no such history (naive communities) over a 4‐yr period in which a major flood disturbance occurred. Comparing communities of identical species composition, we found that selected communities had temporally more stable biomass than naive communities, especially at low species richness. Furthermore, selected communities showed greater biomass recovery after flooding, resulting in more stable post‐flood productivity. In contrast to a previous study, the positive diversity–stability relationship was maintained after the flooding. Our results were consistent across three soil treatments simulating the presence or absence of co‐selected microbial communities. We suggest that prolonged exposure of plant populations to a particular community context and abiotic site conditions can increase ecosystem temporal stability and resilience due to short‐term evolution. A history of co‐occurrence can in part compensate for species loss, as can high plant diversity in part compensate for the missing opportunity of such adaptive adjustments.

The diversity of primary producers strongly affects the structure and diversity of species assemblages at other trophic levels. However, limited knowledge exists of how plant diversity effects at small spatial scales propagate to consumer communities at larger spatial scales. We assessed arthropod community β and γ-diversity in response to experimentally manipulated plant community richness in two long-term grassland biodiversity experiments (Jena, Germany and Cedar Creek, USA) replicated over two years. We calculated arthropod species turnover among all plot combinations (β-diversity), and accumulated number of arthropod species occurring on (1) all pairwise plot combinations and (2) 40 randomly selected six-plot combinations (γ-diversity). The components of arthropod diversity were tested against two measures of plant diversity, namely average plant α-diversity ( PSR ¯ ) and the average difference in plant α-diversity between plots (ΔPSR). Whereas PSR ¯ points to the overall importance of plant α-diversity for arthropod community turnover and diversity on a larger scale, ΔPSR represents the role of habitat heterogeneity. We demonstrate that arthropod γ-diversity is supported by high, homogeneous plant α-diversity, despite lower arthropod β-diversity among high- compared to low-diversity plant communities. We also show that, in six-plot combinations, average plant α-diversity has a positive influence on arthropod γ-diversity only when homogeneity in plant α-diversity is also high. Varying heterogeneity in six-plot combinations showed that combinations consisting solely of plots with an intermediate level of plant α-diversity support a higher number of arthropod species compared to combinations that contain a mix of high- and low-diversity plots. In fact, equal levels of arthropod diversity were found for six-plot combinations with only intermediate or high plant α-diversity, due to saturating benefits of local and larger-scale plant diversity for higher trophic levels. Our results, alongside those of recent observational studies, strongly suggest that maintaining high α-diversity in plant communities is important for conserving multiple components of arthropod diversity. As arthropods carry out a range of essential ecosystem functions, such as pollination and natural pest-control, our findings provide crucial insight for effective planning of human-dominated landscapes to maximize both ecological and economic benefits in grassland systems.

Increasing plant diversity commonly enhances standing biomass and other ecosystem functions (i.e., carbon fluxes, water use efficiency, herbivory). The standing biomass is correlated with vegetation volume, which describes plant biomass allocation within a complex canopy structure. As the canopy structure of plant communities is not static throughout time, it is expected that its changes also control diversity effects on ecosystem functioning. Yet, most studies are based on one or two measures of ecosystem function per year. Here, we examine the temporal effects of diversity of grassland communities on canopy structural components in high temporal (bi-weekly throughout the growing season) and spatial resolutions as a proxy for ecosystem functioning. Using terrestrial laser scanning, we estimate metrics of vertical structure, such as biomass distribution (evenness) and highest biomass allocation (center of gravity) along height strata. For horizontal metrics, we calculated community stand gaps and canopy surface variation. Our findings show that species-rich communities start filling the vertical space (evenness) earlier in the growing season, suggesting a more extended period of resource use (i.e., light-harvesting). Moreover, more diverse communities raised their center of gravity only at the peak of biomass in spring, likely triggered by higher interspecific competition inducing higher biomass allocation at upper layers of the canopy. Furthermore, richer communities were clumpier only after mowing, revealing species-specific differences in regrowth. Lastly, species richness strongly affected canopy variation when the phenology status and height differences were maximal, suggesting differences in plant functional strategies (space to grow, resource use, and flowering phenology). Therefore, the effects of diversity on ecosystem functions depending on those structural components such as biomass production, decomposition, and herbivory, may also change throughout the season due to various mechanisms, such as niche differences, increased complementarity, and temporal and spatial variation in biological activity.

Biodiversity-ecosystem function experiments test how species diversity influences fundamental ecosystem processes. Historically, arthropod driven functions, such as herbivory and pest-control, have been thought to be influenced by direct and indirect associations among species. Although a number of studies have evaluated how plant diversity affects arthropod communities and arthropod-mediated ecosystem processes, it remains unclear whether diversity effects on arthropods are sufficiently consistent over time such that observed responses can be adequately predicted by classical hypotheses based on associational effects. By combining existing results from a long-term grassland biodiversity experiment (Jena Experiment) with new analyses, we evaluate the consistency of consumer responses within and across taxonomic, trophic, and trait-based (i.e. vertical stratification) groupings, and we consider which changes in arthropod community composition are associated with changes in consumer-mediated ecosystem functions. Overall, higher plant species richness supported more diverse and complex arthropod communities and this pattern was consistent across multiple years. Vegetation-associated arthropods responded more strongly to changes in plant species richness than ground-dwelling arthropods. Additionally, increases in plant species richness were associated with shifts in the species-abundance distributions for many, but not all taxa. For example, highly specialized consumers showed a decrease in dominance and an increase in the number of rare species with increasing plant species richness. Most ecosystem processes investigated responded to increases in plant species richness in the same way as the trophic group mediating the process, e.g. both herbivory and herbivore diversity increase with increasing plant species richness. In the Jena Experiment and other studies, inconsistencies between predictions based on classic hypotheses of associational effects and observed relationships between plant species richness and arthropod diversity likely reflect the influence of multi-trophic community dynamics and species functional trait distributions. Future research should focus on testing a broader array of mechanisms to unravel the biological processes underlying the biodiversity-ecosystem functioning relationships.

Biodiversity ensures ecosystem functioning and provisioning of ecosystem services, but it remains unclear how biodiversity-ecosystem multifunctionality relationships depend on the identity and number of functions considered. Here, we demonstrate that ecosystem multifunctionality, based on 82 indicator variables of ecosystem functions in a grassland biodiversity experiment, increases strongly with increasing biodiversity. Analysing subsets of functions showed that the effects of biodiversity on multifunctionality were stronger when more functions were included and that the strength of the biodiversity effects depended on the identity of the functions included. Limits to multifunctionality arose from negative correlations among functions and functions that were not correlated with biodiversity. Our findings underline that the management of ecosystems for the protection of biodiversity cannot be replaced by managing for particular ecosystem functions or services and emphasize the need for specific management to protect biodiversity. More plant species from the experimental pool of 60 species contributed to functioning when more functions were considered. An individual contribution to multifunctionality could be demonstrated for only a fraction of the species.

Changes to primary producer diversity can cascade up to consumers and affect ecosystem processes. Although the effect of producer diversity on higher trophic groups have been studied, these studies often quantify taxonomy-based measures of biodiversity, like species richness, which do not necessarily reflect the functioning of these communities. In this study, we assess how plant species richness affects the functional composition and diversity of higher trophic levels and discuss how this might affect ecosystem processes, such as herbivory, predation and decomposition. Based on six different consumer traits, we examined the functional composition of arthropod communities sampled in experimental plots that differed in plant species richness. The two components we focused on were functional variation in the consumer community structure (functional structure) and functional diversity, expressed as functional richness, evenness and divergence. We found a consistent positive effect of plant species richness on the functional richness of herbivores, carnivores, and omnivores, but not decomposers, and contrasting patterns for functional evenness and divergence. Increasing plant species richness shifted the omnivore community to more predatory and less mobile species, and the herbivore community to more specialized and smaller species. This was accompanied by a shift towards more species occurring in the vegetation than in the ground layer. Our study shows that plant species richness strongly affects the functional structure and diversity of aboveground arthropod communities. The observed shifts in body size (herbivores), specialization (herbivores), and feeding mode (omnivores) together with changes in the functional diversity may underlie previously observed increases in herbivory and predation in plant communities of higher diversity.

The magnitude and frequency of extreme weather events are predicted to increase in the future due to ongoing climate change. In particular, floods and droughts resulting from climate change are thought to alter the ecosystem functions and stability. However, knowledge of the effects of these weather events on soil fauna is scarce, although they are key towards functioning of terrestrial ecosystems. Plant species richness has been shown to affect the stability of ecosystem functions and food webs. Here, we used the occurrence of a natural flood in a biodiversity grassland experiment that was followed by a simulated summer drought experiment, to investigate the interactive effects of plant species richness, a natural flood, and a subsequent summer drought on nematode communities. Three and five months after the natural flooding, effects of flooding severity were still detectable in the belowground system. We found that flooding severity decreased soil nematode food-web structure (loss of K-strategists) and the abundance of plant feeding nematodes. However, high plant species richness maintained higher diversity and abundance of higher trophic levels compared to monocultures throughout the flood. The subsequent summer drought seemed to be of lower importance but reversed negative flooding effects in some cases. This probably occurred because the studied grassland system is well adapted to drought, or because drought conditions alleviated the negative impact of long-term soil waterlogging. Using soil nematodes as indicator taxa, this study suggests that high plant species richness can maintain soil food web complexity after consecutive environmental perturbations.

Recent studies have shown that the diversity of flowering plants can enhance pollinator richness and visitation frequency and thereby increase the resilience of pollination. It is assumed that flower traits explain these effects, but it is still unclear which flower traits are responsible, and knowing that, if pollinator richness and visitation frequency are more driven by mass-ratio effects (mean trait values) or by trait diversity. Here, we analyse a three-year data set of pollinator observations collected in a European grassland plant diversity experiment (The Jena experiment). The data entail comprehensive flower trait measurements, including reward traits (nectar and pollen amount), morphological traits (height, symmetry, area, colour spectra) and chemical traits (nectar-amino acid and nectar-sugar concentration). We test if pollinator species richness and visitation frequency of flower communities depend on overall functional diversity combining all flower traits within a community, single trait diversities (within trait variation) and community-weighted means of the single traits, using Bayesian inference. Overall functional diversity did not affect pollinator species richness, but reduced visitation frequency. When looking at individual flower traits separately, we found that single trait diversity of flower reflectance and flower morphology were important predictors of pollinator visitation frequency. Moreover, independent of total flower abundance, community-weighted means of flower height, area, reflectance, nectar-sugar concentration and nectar-amino acid concentration strongly affected both pollinator species richness and visitation frequency. Our results, challenge the idea that functional diversity always positively affects ecosystem functions. Nonetheless, we demonstrate that both single trait diversity and mass-ratio effects of flower traits play an important role for diverse and frequent flower visits, which underlines the functionality of flower traits for pollination services.

Earth is home to over 350,000 vascular plant species1 that differ in their traits in innumerable ways. Yet, a handful of functional traits can help explaining major differences among species in photosynthetic rate, growth rate, reproductive output and other aspects of plant performance2–6. A key challenge, coined “the Holy Grail” in ecology, is to upscale this understanding in order to predict how natural or anthropogenically driven changes in the identity and diversity of co-occurring plant species drive the functioning of ecosystems7, 8. Here, we analyze the extent to which 42 different ecosystem functions can be predicted by 41 plant traits in 78 experimentally manipulated grassland plots over 10 years. Despite the unprecedented number of traits analyzed, the average percentage of variation in ecosystem functioning that they jointly explained was only moderate (32.6%) within individual years, and even much lower (12.7%) across years. Most other studies linking ecosystem functioning to plant traits analyzed no more than six traits, and when including either only six random or the six most frequently studied traits in our analysis, the average percentage of explained variation in across-year ecosystem functioning dropped to 4.8%. Furthermore, different ecosystem functions were driven by different traits, with on average only 12.2% overlap in significant predictors. Thus, we did not find evidence for the existence of a small set of key traits able to explain variation in multiple ecosystem functions across years. Our results therefore suggest that there are strong limits in the extent to which we can predict the long-term functional consequences of the ongoing, rapid changes in the composition and diversity of plant communities that humanity is currently facing.

Biodiversity often enhances ecosystem functioning likely due to multiple, often temporarily separated drivers. Yet, most studies are based on one or two snapshot measurements per year. We estimated productivity using bi-weekly estimates of high-resolution canopy height in 2 years with terrestrial laser scanning (TLS) in a grassland diversity experiment. We measured how different facets of plant diversity (functional dispersion [FDis], functional identity [PCA species scores], and species richness [SR]) predict aboveground biomass over time. We found strong intra- and inter-annual variability in the relative importance of different mechanisms underlying the diversity effects on mean canopy height, i.e., resource partitioning (via FDis) and identity effects (via species scores), respectively. TLS is a promising tool to quantify community development non-destructively and to unravel the temporal dynamics of biodiversity-ecosystem functioning mechanisms. Our results show that harvesting at estimated peak biomass—as done in most grassland experiments—may miss important variation in underlying mechanisms driving cumulative biomass production.

International audience; Concern about the functional consequences of unprecedented loss in biodiversity has prompted biodiversity-ecosystem functioning (BEF) research to become one of the most active fields of ecological research in the past 25 years. Hundreds of experiments have manipulated biodiversity as an independent variable and found compelling support that the functioning of ecosystems increases with the diversity of their ecological communities. This research has also identified some of the mechanisms underlying BEF relationships, some context-dependencies of the strength of relationships, as well as implications for various ecosystem services that mankind depends upon. In this paper, we argue that a multitrophic perspective of biotic interactions in random and non-random biodiversity change scenarios is key to advance future BEF research and to address some of its most important remaining challenges. We discuss that the study and the quantification of multitrophic interactions in space and time facilitates scaling up from small-scale biodiversity manipulations and ecosystem function assessments to management-relevant spatial scales across ecosystem boundaries. We specifically consider multitrophic conceptual frameworks to understand and predict the context-dependency of BEF relationships. Moreover, we highlight the importance of the eco-evolutionary underpinnings of multitrophic BEF relationships. We outline that FAIR data (meeting the standards of findability, accessibility, interoperability, and reusability) and reproducible processing will be key to advance this field of research by making it more integrative. Finally, we show how these BEF insights may be implemented for ecosystem management, society, and policy. Given that human well-being critically depends on the multiple services provided by diverse, multitrophic communities, integrating the approaches of evolutionary ecology, community ecology, and ecosystem ecology in future BEF research will be key to refine conservation targets and develop sustainable management strategies.

Human activities are causing major changes in biological communities worldwide. Due to concern about the consequences of these changes, an academic conversation about biodiversity and ecosystem functioning (BEF) has emerged over the last few decades. Here we use a keyword co-occurrence analysis to characterize and review 28 years of research focused on these terms. We find that the rapidly growing literature has developed in four research domains. The first two domains “BEF Experiments” and “Science Policy” emerge early, and persist through time, as core research areas with emphases on experiments and management, respectively. The second two domains, “Agricultural Landscapes” and “Aquatic Food Webs”, arise as integrative domains that connect divisions in scientific discussion surrounding BEF Experiments and Science Policy. Terms related to species interactions (i.e. pollinator, predator, food web) appear more commonly in the two integrative domains reflecting shared interests of many scientists focusing on biodiversity and ecosystem functioning. Despite shared interests in food webs, research in the four domains differ with respect to their spatial scale, baseline comparisons, and currency of measurements. Food-web research that bridges these divides should be pushed to the forefront of biodiversity and ecosystem functioning research priorities.

Patterns of feeding interactions between species are thought to influence the stability of communities and the flux of nutrients and energy through ecosystems. However, surprisingly few well-resolved food webs allow us to evaluate factors that influence the architecture of species interactions. We constructed a meta food web consisting of 714 invertebrate species collected over 9 years of suction and pitfall sampling campaigns in the Jena Experiment, a long-term grassland biodiversity experiment located in Jena, Germany. We summarize information on the 51,496 potential trophic links, which were established using information on diet specificity and species traits that typically constrain feeding interactions (trophic group, body size, and vertical stratification). The list of species identities, traits, and link-derivation rules will be useful not only for tests of plant diversity effects on food web structure within the Jena Experiment, but also for considering consistent construction of food webs from empirical data, and for comparisons of network structure across ecosystems. No copyright or proprietary restrictions are associated with the use of this data set other than citation of this Data Paper.

Plant elemental composition can indicate resource limitation, and changes in key elemental ratios (e.g. plant C:N ratios) can influence rates including herbivory, nutrient recycling, and pathogen infection. Although plant stoichiometry can influence ecosystem-level processes, very few studies have addressed whether and how plant C:N stoichiometry changes with plant diversity and composition. Here, using two long-term experimental manipulations of plant diversity (Jena and Cedar Creek), we test whether plant richness (species and functional groups) or composition (functional group proportions) affects temporal trends and variability of community-wide C:N stoichiometry. Site fertility determined the initial community-scale C:N ratio. Communities growing on N-poor soil (Cedar Creek) began with higher C:N ratios than communities growing on N-rich soil (Jena). However, site-level plant C:N ratios converged through time, most rapidly in high diversity plots. In Jena, plant community C:N ratios increased. This temporal trend was stronger with increasing richness. However, temporal variability of C:N decreased as plant richness increased. In contrast, C:N decreased over time at Cedar Creek, most strongly at high species and functional richness, whereas the temporal variability of C:N increased with both measures of diversity at this site. Thus, temporal trends in the mean and variability of C:N were underlain by concordant changes among sites in functional group proportions. In particular, the convergence of community-scale C:N over time at these very different sites was mainly due to increasing proportions of forbs at both sites, replacing high mean C:N (C4 grasses, Cedar Creek) or low C:N (legumes, Jena) species. Diversity amplified this convergence; although temporal trends differed in sign between the sites, these trends increased in magnitude with increasing species richness. Our results suggest a predictive mechanistic link between trends in plant diversity and functional group composition and trends in the many ecosystem rates that depend on aboveground community C:N. Synthesis We compared the effect of plant diversity on the temporal dynamics of community stoichiometry in two long-term grassland diversity experiments: the Cedar Creek and Jena Experiments. Changes in community C:N ratios were accelerated by increasing diversity at both sites, but in opposite directions depending on soil fertility. Stoichiometry changes were driven by shifts of functional group composition differing in their elemental compositions, the identity of the functional groups depending on the site. Thus, we highlighted that community turnover constrained the effect of diversity on plant stoichiometry at both sites.

Plant diversity is an important driver of nitrogen and phosphorus stocks in aboveground plant biomass of grassland ecosystems, but plant diversity effects on other elements also important for plant growth are less understood. We tested whether plant species richness, functional group richness or the presence/absence of particular plant functional groups influences the Si and Ca concentrations (mmol g(-1)) and stocks (mmol m(-2)) in aboveground plant biomass in a large grassland biodiversity experiment (Jena Experiment). In the experiment including 60 temperate grassland species, plant diversity was manipulated as sown species richness (1, 2, 4, 8, 16) and richness and identity of plant functional groups (1-4; grasses, small herbs, tall herbs, legumes). We found positive species richness effects on Si as well as Ca stocks that were attributable to increased biomass production. The presence of particular functional groups was the most important factor explaining variation in aboveground Si and Ca stocks (mmol m(-2)). Grass presence increased the Si stocks by 140 % and legume presence increased the Ca stock by 230 %. Both the presence of specific plant functional groups and species diversity altered Si and Ca stocks, whereas Si and Ca concentration were affected mostly by the presence of specific plant functional groups. However, we found a negative effect of species diversity on Si and Ca accumulation, by calculating the deviation between mixtures and mixture biomass proportions, but in monoculture concentrations. These changes may in turn affect ecosystem processes such as plant litter decomposition and nutrient cycling in grasslands.

Understanding factors that increase ecosystem stability is critical in the face of environmental change. Experiments simulating species loss from grassland ecosystems have shown that losing biodiversity decreases the ability of ecosystems to buffer negative effects of disturbances. However, as the originally sown experimental communities with reduced biodiversity develop, plant evolutionary processes or the assembly of interacting soil organisms may allow them to develop stability and resilience over time. We explored such effects in a long-term grassland biodiversity experiment with plant communities with either a history of co-occurrence (selected communities) or no such history (naïve communities) over a four-year period in which a major flood disturbance occurred.We found selected communities had temporally more stable biomass than the same communities of naïve plants, especially at low species richness. Furthermore, selected communities showed greater short-term biomass recovery after flooding, resulting in more stable post-flood productivity. In contrast to a previous study, the positive diversity–stability relationship was maintained after the flooding. Our results were consistent across three soil treatments simulating the presence or absence of co-selected microbial communities. We suggest that prolonged exposure of plant populations to a particular community context and abiotic site conditions can increase ecosystem temporal stability and resistance to disturbance. We argue that selection during the course of a biodiversity experiment is the most parsimonious explanation for these effects. A history of co-occurrence can in part compensate for species loss, as can high plant diversity in part compensate for the missing opportunity of such adaptive adjustments.

Plant species richness (PSR) increases nutrient uptake which depletes bioavailable nutrient pools in soil. No such relationship between plant uptake and availability in soil was found for phosphorus (P). We explored PSR effects on P mobilisation [phosphatase activity (PA)] in soil. PA increased with PSR. The positive PSR effect was not solely due to an increase in Corg concentrations because PSR remained significant if related to PA:Corg . An increase in PA per unit Corg increases the probability of the temporal and spatial match between substrate, enzyme and microorganism potentially serving as an adaption to competition. Carbon use efficiency of microorganisms (Cmic :Corg ) increased with increasing PSR while enzyme exudation efficiency (PA:Cmic ) remained constant. These findings suggest the need for efficient C rather than P cycling underlying the relationship between PSR and PA. Our results indicate that the coupling between C and P cycling in soil becomes tighter with increasing PSR.

Plant functional characteristics may drive plant species richness effects on ecosystem processes. Consequently, the focus of biodiversity–ecosystem functioning (BEF) experiments has expanded from the manipulation of plant species richness to manipulating functional trait composition. Involving ecophysiological plant traits in the experimental design might allow for a better understanding of how species loss alters ecosystem processes. Here we provide the theoretical background, design and first results of the ‘Trait-Based Biodiversity Experiment’ (TBE), established in 2010 that directly manipulates the trait composition of experimental plant communities. Analysis of six plant traits related to resource acquisition and use were analyzed using principal component analysis of 60 grassland species. The resulting two main axes describe gradients in functional similarity, and were used as the basis for designing plant communities with different functional and species diversity levels. Using such an approach allowed us to manipulate different levels of complementarity in spatial and temporal plant resource acquisition. In contrast to previous biodiversity experiments, the TBE is designed according to more realistic scenarios of non-random species loss along orthogonal axes of species trait dissimilarities. This allows us to tease apart the relative importance of selection and complementarity effects on multiple ecosystem processes, and to mechanistically study the consequences of plant community simplification.

Plant diversity is a key driver of ecosystem functioning best documented for its influence on plant productivity. The strength and direction of plant diversity effects on species interactions across trophic levels are less clear. For example, with respect to the interactions between herbivorous invertebrates and plants, a number of competing hypotheses have been proposed that predict either increasing or decreasing community herbivory with increasing plant species richness. We investigated foliar herbivory rates and consumed leaf biomass along an experimental grassland plant diversity gradient in year eight after establishment. The gradient ranged from one to 60 plant species and manipulated also functional group richness (from one to four functional groups-legumes, grasses, small herbs, and tall herbs) and plant community composition. Measurements in monocultures of each plant species showed that functional groups differed in the quantity and quality of herbivory damage they experienced, with legumes being more damaged than grasses or non-legume herbs. In mixed plant communities, herbivory increased with plant diversity and the presence of two key plant functional groups in mixtures had a positive (legumes) and a negative (grasses) effect on levels of herbivory. Further, plant community biomass had a strong positive impact on consumed leaf biomass, but little effect on herbivory rates. Our results contribute detailed data from a well-established biodiversity experiment to a growing body of evidence suggesting that an increase of herbivory with increasing plant diversity is the rule rather than an exception. Considering documented effects of herbivory on other ecosystem functions and the increase of herbivory with plant diversity, levels of herbivory damage might not only be a result, but also a trigger within the diversity-productivity relationship.

In many environments land use intensification is likely to result in a decrease in species richness and in an increase in eutrophication. Although the importance of both factors for higher trophic levels such as insect herbivores is well documented, their impact has rarely been studied in combination. Herbivorous insects have a strong impact on the functioning of ecosystems and it is therefore important to understand how they are affected by eutrophication in high or low diversity environments. We used a grassland biodiversity experiment to investigate the combined effect of fertilization and plant diversity loss on the fitness of the generalist grasshopper Chorthippus parallelus by rearing grasshopper nymphs for four weeks in cages on unfertilized or fertilized (NPK) subplots across a species richness gradient from 1 to 60 plant species. Survival, the number of oothecae, body mass and the number of hatchlings were measured separately for each cage. Plant diversity had no effect on any of the grasshopper fitness measures, neither in unfertilized nor in fertilized plots. NPK-fertilization reduced grasshopper survival but increased body mass of males and reproductive success of the surviving females. Fertilization effects were not mediated by plant community structure, productivity or composition, suggesting that higher food plant quality was one of the main drivers. There was no interaction between plant diversity and fertilization on any of the measures. In conclusion, an increase in eutrophication, in both species-rich and species-poor grasslands, could lead to higher reproductive success and therefore higher abundances of herbivorous insects including insect pests, with fertilization effects dominating plant diversity effects.

In the past two decades, a large number of studies have investigated the relationship between biodiversity and ecosystem functioning, most of which focussed on a limited set of ecosystem variables. The Jena Experiment was set up in 2002 to investigate the effects of plant diversity on element cycling and trophic interactions, using a multi-disciplinary approach. Here, we review the results of 15 years of research in the Jena Experiment, focussing on the effects of manipulating plant species richness and plant functional richness. With more than 85,000 measures taken from the plant diversity plots, the Jena Experiment has allowed answering fundamental questions important for functional biodiversity research.[br/] First, the question was how general the effect of plant species richness is, regarding the many different processes that take place in an ecosystem. About 45% of different types of ecosystem processes measured in the 'main experiment', where plant species richness ranged from 1 to 60 species, were significantly affected by plant species richness, providing strong support for the view that biodiversity is a significant driver of ecosystem functioning. Many measures were not saturating at the 60-species level, but increased linearly with the logarithm of species richness. There was, however, great variability in the strength of response among different processes. One striking pattern was that many processes, in particular belowground processes, took several years to respond to the manipulation of plant species richness, showing that biodiversity experiments have to be long-term, to distinguish trends from transitory patterns. In addition, the results from the Jena Experiment provide further evidence that diversity begets stability, for example stability against invasion of plant species, but unexpectedly some results also suggested the opposite, e.g. when plant communities experience severe perturbations or elevated resource availability. This highlights the need to revisit diversity-stability theory.[br/] Second, we explored whether individual plant species or individual plant functional groups, or biodiversity itself is more important for ecosystem functioning, in particular biomass production. We found strong effects of individual species and plant functional groups on biomass production, yet these effects mostly occurred in addition to, but not instead of, effects of plant species richness.[br/] Third, the Jena Experiment assessed the effect of diversity on multitrophic interactions. The diversity of most organisms responded positively to increases in plant species richness, and the effect was stronger for above-than for belowground organisms, and stronger for herbivores than for carnivores or detritivores. Thus, diversity begets diversity. In addition, the effect on organismic diversity was stronger than the effect on species abundances.[br/] Fourth, the Jena Experiment aimed to assess the effect of diversity on N, P and C cycling and the water balance of the plots, separating between element input into the ecosystem, element turnover, element stocks, and output from the ecosystem. While inputs were generally less affected by plant species richness, measures of element stocks, turnover and output were often positively affected by plant diversity, e.g. carbon storage strongly increased with increasing plant species richness. Variables of the N cycle responded less strongly to plant species richness than variables of the C cycle.[br/] Fifth, plant traits are often used to unravel mechanisms underlying the biodiversity-ecosystem functioning relationship. In the Jena Experiment, most investigated plant traits, both above-and belowground, were plastic and trait expression depended on plant diversity in a complex way, suggesting limitation to using database traits for linking plant traits to particular functions.[br/] Sixth, plant diversity effects on ecosystem processes are often caused by plant diversity effects on species interactions. Analyses in the Jena Experiment including structural equation modelling suggest complex interactions that changed with diversity, e.g. soil carbon storage and greenhouse gas emission were affected by changes in the composition and activity of the belowground microbial community. Manipulation experiments, in which particular organisms, e.g. belowground invertebrates, were excluded from plots in split-plot experiments, supported the important role of the biotic component for element and water fluxes.[br/] Seventh, the Jena Experiment aimed to put the results into the context of agricultural practices in managed grasslands. The effect of increasing plant species richness from 1 to 16 species on plant biomass was, in absolute terms, as strong as the effect of a more intensive grassland management, using fertiliser and increasing mowing frequency. Potential bioenergy production from high-diversity plots was similar to that of conventionally used energy crops. These results suggest that diverse 'High Nature Value Grasslands' are multifunctional and can deliver a range of ecosystem services including production-related services.[br/] A final task was to assess the importance of potential artefacts in biodiversity-ecosystem functioning relationships, caused by the weeding of the plant community to maintain plant species composition. While the effort ( in hours) needed to weed a plot was often negatively related to plant species richness, species richness still affected the majority of ecosystem variables. Weeding also did not negatively affect monoculture performance; rather, monocultures deteriorated over time for a number of biological reasons, as shown in plant-soil feedback experiments.[br/] To summarize, the Jena Experiment has allowed for a comprehensive analysis of the functional role of biodiversity in an ecosystem. A main challenge for future biodiversity research is to increase our mechanistic understanding of why the magnitude of biodiversity effects differs among processes and contexts. It is likely that there will be no simple answer. For example, among the multitude of mechanisms suggested to underlie the positive plant species richness effect on biomass, some have received limited support in the Jena Experiment, such as vertical root niche partitioning. However, others could not be rejected in targeted analyses. Thus, from the current results in the Jena Experiment, it seems likely that the positive biodiversity effect results from several mechanisms acting simultaneously in more diverse communities, such as reduced pathogen attack, the presence of more plant growth promoting organisms, less seed limitation, and increased trait differences leading to complementarity in resource uptake. Distinguishing between different mechanisms requires careful testing of competing hypotheses. Biodiversity research has matured such that predictive approaches testing particular mechanisms are now possible.

Pollinators play a key role in the reproduction of most plant species, and pollinator and plant diversity are often related. We studied an experimental gradient of plant species richness for a better understanding of plantpollinator community interactions and their temporal variability, because in non-experimental field surveys plant richness is often confounded with gradients in management, soil fertility, and community composition. We observed pollinator species richness and frequency of visits six times in 73 plots over two years, and used advanced statistical analysis to account for the high number of zeroes that often occur in count data of rare species. The frequency of pollinator visits increased linearly with both the blossom cover and the number of flowering plant species, which was closely related to the total number of plant species, whereas the number of pollinator species followed a saturation curve. The presence of particularly attractive plant species was only important for the frequency of flower visits, but not to the richness of pollinators. Plant species richness, blossom cover, and the presence of attractive plant species enhanced the temporal stability in the frequency of pollinator visits. In conclusion, grasslands with high plant diversity enhance and stabilize frequent and diverse flower visitations, which should sustain effective pollination and plant reproduction.

Climate change is expected to increase the frequency and magnitude of extreme weather events. It is therefore of major importance to identify the community attributes that confer stability in ecological communities during such events. In June 2013, a flood event affected a plant diversity experiment in Central Europe (Jena, Germany). We assessed the effects of plant species richness, functional diversity, flooding intensity and community means of functional traits on different measures of stability (resistance, resilience and raw biomass changes from pre-flood conditions). Surprisingly, plant species richness reduced community resistance in response to the flood. This was mostly because more diverse communities grew more immediately following the flood. Raw biomass increased over the previous year; this resulted in decreased absolute value measures of resistance. There was no clear response pattern for resilience. We found that functional traits drove these changes in raw biomass: communities with a high proportion of late-season, short-statured plants with dense, shallow roots and small leaves grew more following the flood. Late-growing species probably avoided the flood, whereas greater root length density might have allowed species to better access soil resources brought from the flood, thus growing more in the aftermath. We conclude that resource inputs following mild floods may favour the importance of traits related to resource acquisition and be less associated with flooding tolerance.

In a grassland plant diversity experiment, we studied if plant species richness induces transgressive overyielding in soil microbial biomass, the definition of which is greater microbial biomass with a plant mixture than in each of the plant monocultures. In June and November of 2012 soil microbial biomass increased significantly with plant species richness. Overyielding occurred in more than 61% of 24 plots containing six species, and transgressive overyielding occurred in 13–21% of the plots depending on season. Plots with nine species showed overyielding in all cases, and 25–50% showed transgressive overyielding. Transgressive underyielding occurred in a few plots with six species in June. Our results indicate plant complementarity effects on soil microbial biomass in diverse plant communities across seasons and stress the relevance of simplification of plant communities for soil processes.

Identifying patients in cardiology who are able to benefit from more detailed psychosocial diagnostics or interventions is still problematic in every day hospital care. Existing psychosocial screening instruments--either self-evaluation questionnaires and evaluation interviews carried out by doctors--are discussed. A new screening instrument for evaluating patients in cardiology is presented: The Luebeck Interview for Psychosocial Screening (LIPS). With help of LIPS patients are evaluated by their doctor through a short semistructured interview referring to the patients' psychosocial situation. Test statistical data of the instrument gained through 194 patients are presented. For a subgroup of 80 patients the validity for predicting their quality of life six weeks after discharge form hospital by psychosocial assessment in hospital is presented. Data on reliability and validity of LIPS appear to range from acceptable to good. Predicting quality of life by self-report inventories reaches a better fit than a prediction by LIPS as an evaluation from the doctor's perspective. The errors of the prediction by both methods are highly correlated. Further research on LIPS combined with a structured clinical Interview for mental disorders seems to be desirable.

Ecosystem responses to changes in species diversity are often studied individually. However, changes in species diversity can simultaneously influence multiple interdependent ecosystem functions. Therefore, an important challenge is to determine when and how changes in species diversity that influence one function will also drive changes in other functions. By providing the underlying structure of species interactions, ecological networks can quantify connections between biodiversity and multiple ecosystem functions. Here, we review parallels in the conceptual development of biodiversity–ecosystem functioning (BEF) and food web theory (FWT) research. Subsequently, we evaluate three common principles that unite these two research areas by explaining the patterns, concentrations, and direction of the flux of nutrients and energy through the species in diverse interaction webs. We give examples of combined BEF–FWT approaches that can be used to identify vulnerable species and habitats and to evaluate links that drive trade-offs between multiple ecosystems functions. These combined approaches reflect promising trends towards better management of biodiversity in landscapes that provide essential ecosystem services supporting human well-being.

Biodiversity experiments generally report a positive effect of plant biodiversity on aboveground biomass (overyielding), which typically increases with time. Various studies also found overyielding for belowground plant biomass, but this has never been measured over time. Also, potential underlying mechanisms have remained unclear. Differentiation in rooting patterns among plant species and plant functional groups has been proposed as a main driver of the observed biodiversity effect on belowground biomass, leading to more efficient belowground resource use with increasing diversity, but so far there is little evidence to support this. We analyzed standing root biomass and its distribution over the soil profile, along a 1-16 species richness gradient over eight years in the Jena Experiment in Germany, and compared belowground to aboveground overyielding. In our long-term dataset, total root biomass increased with increasing species richness but this effect was only apparent after four years. The increasingly positive relationship between species richness and root biomass, explaining 12% of overall variation and up to 28% in the last year of our study, was mainly due to decreasing root biomass at low diversity over time. Functional group composition strongly affected total standing root biomass, explaining 44% of variation, with grasses and legumes having strong overall positive and negative effects, respectively. Functional group richness or interactions between functional group presences did not strongly contribute to overyielding. We found no support for the hypothesis that vertical root differentiation increases with species richness, with functional group richness or composition. Other explanations, such as stronger negative plant-soil feedbacks in low-diverse plant communities on standing root biomass and vertical distribution should be considered.