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Drought is a major stressor to soil microbial communities, and the intensification of climate change is predicted to increase hydric stress worldwide in the coming decades. As a possible mitigating factor for the consequences of prolonged drought periods, above and belowground biodiversity can increase ecosystem resistance and resilience by improving metabolic redundancy and complementarity as biodiversity increases. Here, we investigated the interaction effect between plant richness and successive, simulated summer drought on soil microbial communities during a period of 9 years.To do that, we made use of a well-established biodiversity experiment (The Jena Experiment) to investigate the response of microbial richness and community composition to successive drought periods alongside a plant richness gradient, which covers 1-, 2-, 4-, 8-, 16-, and 60-species plant communities. Plots were covered from natural precipitation by installing rain shelters 6 weeks every summer. Bulk soil samples were collected 1 year after the last summer drought was simulated. Our data indicate that bacterial richness increased after successive exposure to drought, with the increase being stable along the plant richness gradient. We identified a significant effect of plant species richness on the soil microbial community composition and determined the taxa significantly impacted by drought at each plant richness level. Our data successfully demonstrates that summer drought might have a legacy effect on soil bacterial communities. [ABSTRACT FROM AUTHOR]

Soil is the basis for life on Earth as we know it. Healthy and stable soil is a prerequisite for well-functioning terrestrial ecosystems and has, thus, been proposed to play a key role in plant diversity–ecosystem functioning relationships. The overall objective of this sub-project is to study multidimensional soil stability as affected by plant diversity in a long-term plant diversity experiment. We designed three coordinated work packages (WPs) to comprehensively assess soil multistability to environmental fluctuations and climate extremes by considering the biological, chemical and physical dimensions that are key for soil functioning. We will use all unique facilities and approaches of the Jena Experiment Research Unit by combining synthesis of long-term data in the Main Experiment and the ΔBEF Experiment with performing new soil analyses in the DrY Experiment, the ResCUE Experiment and a joint CoMic Experiment, to gain a better mechanistic understanding of plant diversity–ecosystem functioning relationships. In close collaboration with other sub-projects, we will assess biological, chemical and physical soil properties and stability indicators that will be used to calculate soil multifunctionality and multistability indices. In WP1, we will build on three unique datasets to explore short-term and long-term effects of plant diversity on the stability of soil (microbial) properties. In WP2, we will combine different datasets and approaches to explore if plant diversity effects on the magnitude and stability of soil properties increase with abiotic and biotic stresses. In WP3, we will combine measurements of the above-mentioned dimensions of soil stability to explore if plant diversity increases the stability of multiple soil properties under hot drought. This sub-project is at the heart of the Research Unit by testing the overarching hypotheses outlined in the Coordination Proposal of the Jena Experiment, contributing to all main experiments, sharing data and performing joint sampling campaigns with all sub-projects and, at the same time, introducing a novel concept of soil multistability as affected by plant diversity and climate extremes. We propose to use a combination of simple, high-throughput (e.g. bait-lamina test) and more sophisticated methods (e.g. extracellular polymeric substances analyses) to be able to investigate temporal dynamics of soil processes and their mechanistic basis. Taken together, the results of the three WPs will provide new insights into the stabilising mechanisms of soil properties in the long term and in relation to climate extremes through plant diversity. [ABSTRACT FROM AUTHOR]

It is well known that biodiversity positively affects ecosystem functioning, leading to enhanced ecosystem stability. However, this knowledge is mainly based on analyses using single ecosystem functions, while studies focusing on the stability of ecosystem multifunctionality (EMF) are rare. Taking advantage of a long‐term grassland biodiversity experiment, we studied the effect of plant diversity (1–60 species) on EMF over 5 years, its temporal stability, as well as multifunctional resistance and resilience to a 2‐year drought event. Using split‐plot treatments, we further tested whether a shared history of plants and soil influences the studied relationships. We calculated EMF based on functions related to plants and higher‐trophic levels. Plant diversity enhanced EMF in all studied years, and this effect strengthened over the study period. Moreover, plant diversity increased the temporal stability of EMF and fostered resistance to reoccurring drought events. Old plant communities with shared plant and soil history showed a stronger plant diversity–multifunctionality relationship and higher temporal stability of EMF than younger communities without shared histories. Our results highlight the importance of old and biodiverse plant communities for EMF and its stability to extreme climate events in a world increasingly threatened by global change. [ABSTRACT FROM AUTHOR]

Artificial light at night (ALAN) affects many areas of the world and is increasing globally. To date, there has been limited and inconsistent evidence regarding the consequences of ALAN for plant communities, as well as for the fitness of their constituent species. ALAN could be beneficial for plants as they need light as energy source, but they also need darkness for regeneration and growth. We created model communities composed of 16 plant species sown, exposed to a gradient of ALAN ranging from 'moonlight only' to conditions like situations typically found directly underneath a streetlamp. We measured plant community composition and its production (biomass), as well as functional traits of three plant species from different functional groups (grasses, herbs, legumes) in two separate harvests. We found that biomass was reduced by 33% in the highest ALAN treatment compared to the control, Shannon diversity decreased by 43% and evenness by 34% in the first harvest. Some species failed to establish in the second harvest. Specific leaf area, leaf dry matter content and leaf hairiness responded to ALAN. These responses suggest that plant communities will be sensitive to increasing ALAN, and they flag a need for plant conservation activities that consider impending ALAN scenarios. This article is part of the theme issue 'Light pollution in complex ecological systems'. [ABSTRACT FROM AUTHOR]

Within biodiversity‐ecosystem function research, a major outstanding question is how herbivory, a critical ecosystem function at the base of the food web, changes along gradients of plant biodiversity.Neighbourhood‐level associational effects are hypothesised to be a strong driver of biodiversity‐herbivory relationships, but we lack a successful framework that explains the wide variation observed in the sign and magnitude of plant‐herbivore associational effects.In this study, we combine measurements from a tree biodiversity field experiment with simulation to provide a framework for explaining variation in plant‐herbivore associational effects, particularly when herbivores that feed on many different species (e.g. generalists) cause most damage. We show that monoculture herbivory levels of focal species and their neighbours predict the direction and strength of associational effects. We provide evidence that this may be due to a "spillover effect", in which some insect herbivores attracted to focal individuals ultimately end up feeding on neighbouring individuals.With an empirically parameterised simulation, we explain how spatial organisation modifies biodiversity‐ecosystem function relationships when associational effects operate. We suggest a set of experiments to test the generality of our conceptual framework, to elucidate the underlying mechanisms that produce the patterns we find, and to ultimately increase the predictability of plant‐herbivore associational effects. We conclude by discussing how our results might inform pest management in diversified agroecosystems and reforestation sites.Synthesis. Our results provide a potential framework for explaining why positive and negative plant‐herbivore associational effects are often balanced in systems with primarily generalist herbivores and point to a path forward for predicting when increased plant biodiversity will be associated with increased, decreased or unchanged levels of insect herbivory on individual plant species in such systems. [ABSTRACT FROM AUTHOR]

Plant and soil biodiversity can have significant effects on herbivore resistance mediated by plant metabolites. Here, we disentangled the independent effects of plant diversity and soil legacy on constitutive and herbivore‐induced plant metabolomes of three plant species in two complementary microcosm experiments. First, we grew plants in sterile soil with three different plant diversity levels. Second, single plant species were grown on soil with different plant diversity‐induced soil legacies. We infested a subset of all plants with Spodoptera exigua larvae, a generalist leaf‐chewing herbivore, and assessed foliar and root metabolomes. Neither plant diversity nor soil legacy had significant effects on overall foliar, root, or herbivore‐induced metabolome composition. Herbivore‐induced metabolomes, however, differed from those of control plants. We detected 139 significantly regulated metabolites by comparing plants grown in monocultures with conspecifics growing in plant or soil legacy mixtures. Moreover, plant–plant and plant–soil interactions regulated 141 metabolites in herbivore‐induced plants. Taken together, plant diversity and soil legacy independently alter the concentration and induction of plant metabolites, thus affecting the plant's defensive capability. This is a first step toward disentangling plant and soil biodiversity effects on herbivore resistance, thereby improving our understanding of the mechanisms that govern ecosystem functioning. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Soils are important for ecosystem functioning and service provisioning. Soil communities and their functions, in turn, are strongly promoted by plant diversity, and such positive effects strengthen with time. However, plant diversity effects on soil organic matter have mostly been investigated in the topsoil, and there are only very few long‐term studies. Thus, it remains unclear if plant diversity effects strengthen with time and to which depth these effects extend. Here, we repeatedly sampled soil to 1 m depth in a long‐term grassland biodiversity experiment. We investigated how plant diversity impacted soil organic carbon and nitrogen concentrations and stocks and their stable isotopes 13C and 15N, as well as how these effects changed after 5, 10, and 14 years. We found that higher plant diversity increased carbon and nitrogen storage in the topsoil since the establishment of the experiment. Stable isotopes revealed that these increases were associated with new plant‐derived inputs, resulting in less processed and less decomposed soil organic matter. In subsoils, mainly the presence of specific plant functional groups drove organic matter dynamics. For example, the presence of deep‐rooting tall herbs decreased carbon concentrations, most probably through stimulating soil organic matter decomposition. Moreover, plant diversity effects on soil organic matter became stronger in topsoil over time and reached subsoil layers, while the effects of specific plant functional groups in subsoil progressively diminished over time. Our results indicate that after changing the soil system the pathways of organic matter transfer to the subsoil need time to establish. In our grassland system, organic matter storage in subsoils was driven by the redistribution of already stored soil organic matter from the topsoil to deeper soil layers, for example, via bioturbation or dissolved organic matter. Therefore, managing plant diversity may, thus, have significant implications for subsoil carbon storage and other critical ecosystem services. [ABSTRACT FROM AUTHOR]

Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions. The insurance hypothesis posits that more diverse communities are more stable through time. Here, the authors show that plant biodiversity reduces the spatial variability of productivity in grassland communities, demonstrating that the insurance hypothesis applies also across space. [ABSTRACT FROM AUTHOR]

Positive plant diversity–productivity relationships are known to be driven by complementary resource use via differences in plant functional traits. Moreover, soil properties related to nutrient availability were shown to change with plant diversity over time; however, it is not well‐understood whether and how such plant diversity‐dependent soil changes and associated changes in functional traits contribute to positive diversity–productivity relationships in the long run. To test this, we investigated plant communities of different species richness (1, 2, 6, and 9 species) in a 15‐year‐old grassland biodiversity experiment. We determined community biomass production and biodiversity effects (net biodiversity [NEs], complementarity [CEs], and selection effects [SEs]), as well as community means of plant functional traits and soil properties. First, we tested how these variables changed along the plant diversity gradient and were related to each other. Then, we tested for direct and indirect effects of plant and soil variables influencing community biomass production and biodiversity effects. Community biomass production, NEs, CEs, SEs, plant height, root length density (RLD), and all soil property variables changed with plant diversity and the presence of the dominant grass species Arrhenatherum elatius (increase except for soil pH, which decreased). Plant height and RLD for plant functional traits, and soil pH and organic carbon concentration for soil properties, were the variables with the strongest influence on biomass production and biodiversity effects. Our results suggest that plant species richness and the presence of the dominant species, A. elatius, cause soil organic carbon to increase and soil pH to decrease over time, which increases nutrient availability favoring species with tall growth and dense root systems, resulting in higher biomass production in species‐rich communities. Here, we present an additional process that contributes to the strengthening positive diversity–productivity relationship, which may play a role alongside the widespread plant functional trait‐based explanation. [ABSTRACT FROM AUTHOR]

Insect herbivory is a key process in ecosystem functioning. While theory predicts that plant diversity modulates herbivory, the mechanistic links remain unclear. We postulated that the plant metabolome mechanistically links plant diversity and herbivory.In late summer and in spring, we assessed individual plant above‐ground herbivory rates and metabolomes of seven plant species in experimental plant communities varying in plant species diversity and resource acquisition strategies. In the same communities, we also measured plant individual biomass as well as soil microbial and nematode community composition.Herbivory rates decreased with increasing plant species richness. Path modelling revealed that plant species richness and community resource acquisition strategy correlated with soil community composition. In particular, changes in nematode community composition were related to plant metabolome composition and thereby herbivory rates.Synthesis. These results suggest that soil community composition plays an important role in reducing herbivory rates with increasing plant diversity by changing plant metabolomes. [ABSTRACT FROM AUTHOR]

Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting. [ABSTRACT FROM AUTHOR]

Resources can affect plant productivity and biodiversity simultaneously and thus are key drivers of their relationships in addition to plant–plant interactions. However, most previous studies only focused on a single resource while neglecting the nature of resource multidimensionality. Here we integrated four essential resources for plant growth into a single metric of resource diversity (RD) to investigate its effects on the productivity–biodiversity relationship (PBR) across Chinese grasslands. Results showed that habitats differing in RD have different PBRs—positive in low-resource habitats, but neutral in medium- and high-resource ones—while collectively, a weak positive PBR was observed. However, when excluding direct effects of RD on productivity and biodiversity, the PBR in high-resource habitats became negative, which leads to a unimodal instead of a positive PBR along the RD gradient. By integrating resource effects and changing plant–plant interactions into a unified framework with the RD gradient, our work contributes to uncovering underlying mechanisms for inconsistent PBRs at large scales. [ABSTRACT FROM AUTHOR]

1. Understanding how ecosystem multifunctionality is maintained in naturally assembled communities is crucial, because human activities benefit from multiple functions and services of various ecosystems. However, the effects of aboveand below-ground biodiversity on ecosystem multifunctionality in alpine and boreal moorland ecosystems remain unclear despite their potential as global carbon sinks. 2. Here we evaluated how ecosystem multifunctionality related to primary production and carbon sequestration, which are crucial for global climate regulation, is maintained in natural systems. We disentangled the relationships between diversity and composition of plants and soil microbes (fungi and bacteria) and ecosystem multifunctionality in subalpine moorlands in northern Japan. 3. We found that microbial composition primarily regulated carbon sequestration, whereas plant taxonomic and functional composition were related to all functions considered. Plant and microbial α diversity (diversity within local communities) were not generally related to any single function, highlighting the important roles of specific plant and microbial taxa in determining ecosystem functioning. When single functions were aggregated to ecosystem multifunctionality within local communities, plant and microbial community composition rather than diversity regulated ecosystem multifunctionality. We further found that plant and bacterial taxonomic β diversity (taxonomic turnover between local communities) primarily regulated the dissimilarity of ecosystem multifunctionality between local communities. 4. Synthesis. We provide observational evidence that plant and microbial community composition rather than diversity are essential for sustaining subalpine moorland multifunctionality. Furthermore, plant and bacterial β diversity enhance the dissimilarity of moorland multifunctionality. Our study provides novel insights into biodiversity–ecosystem multifunctionality relationships occurring in nature, and helps to sustain desirable ecosystem functioning to human society. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Diversity loss has been shown to change the soil community; however, little is known about long-term consequences and underlying mechanisms. Here, we investigated how nematode communities are affected by plant species richness and whether this is driven by resource quantity or quality in 15-year-old plant communities of a long-term grassland biodiversity experiment. We extracted nematodes from 93 experimental plots differing in plant species richness, and measured above- and belowground plant biomass production and soil organic carbon concentrations (Corg) as proxies for resource quantity, as well as C/Nleaf ratio and specific root length (SRL) as proxies for resource quality. We found that nematode community composition and diversity significantly differed among plant species richness levels. This was mostly due to positive plant diversity effects on the abundance and genus richness of bacterial-feeding, omnivorous, and predatory nematodes, which benefited from higher shoot mass and soil Corg in species-rich plant communities, suggesting control via resource quantity. In contrast, plant-feeding nematodes were negatively influenced by shoot mass, probably due to higher top–down control by predators, and were positively related to SRL and C/Nleaf, indicating control via resource quality. The decrease of the grazing pressure ratio (plant feeders per root mass) with plant species richness indicated a higher accumulation of plant-feeding nematodes in species-poor plant communities. Our results, therefore, support the hypothesis that soil-borne pathogens accumulate in low-diversity communities over time, while soil mutualists (bacterial-feeding, omnivorous, predatory nematodes) increase in abundance and richness in high-diversity plant communities, which may contribute to the widely-observed positive plant diversity–productivity relationship. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Identifying to what degree inherent characteristics of plant species and their variation in response to their environment regulate the temporal stability of plant populations is important to understand patterns of species coexistence and the stability of ecosystems. Longevity is a key characteristic of plant life history and an important component of demographic storage, but age is usually unknown for herbaceous species.In a 12‐year‐old biodiversity experiment (Jena Experiment) comprising 80 grassland communities with six levels of plant species richness (1, 2, 4, 8, 16 and 60 species) and four levels of functional groups richness (1, 2, 3 and 4 functional groups), we studied populations of 38 dicotyledonous forb species (N = 1,683 plant individuals). The sampled individuals represented three plant functional groups (legumes, small herbs and tall herbs) and two different growth forms (species with long‐lived primary roots and clonal species with rhizomes/stolons). We assessed the age of plant individuals by means of growth ring analysis and related the age of plant populations to their temporal stability in terms of peak biomass production.On average, plant species richness did not affect the mean age of the populations or the maximum age of individuals found in a population. Age of herbs with taproots increased and age of herbs with clonal growth decreased with increasing species richness, cancelling out each other when growth forms were analysed together. Mean population age was lowest for small herbs and highest for tall herbs, while legumes had an intermediate population age. Herbs with a taproot were on average older than herbs with a rhizome. Across all species‐richness levels, populations with older individuals were more stable in terms of biomass production over time.Synthesis. Our study shows for the first time across multiple species that the longevity of forbs is affected by the diversity of the surrounding plant community, and that plant longevity as an important component of demographic storage increases the temporal stability of populations of grassland forb species. [ABSTRACT FROM AUTHOR]

Long‐term biodiversity experiments have shown increasing strengths of biodiversity effects on plant productivity over time. However, little is known about rapid evolutionary processes in response to plant community diversity, which could contribute to explaining the strengthening positive relationship. To address this issue, we performed a transplant experiment with offspring of seeds collected from four grass species in a 14‐year‐old biodiversity experiment (Jena Experiment). We used two‐ and six‐species communities and removed the vegetation of the study plots to exclude plant–plant interactions. In a reciprocal design, we transplanted five "home" phytometers (same origin and actual environment), five "away‐same" phytometers (same species richness of origin and actual environment, but different plant composition), and five "away‐different" phytometers (different species richness of origin and actual environment) of the same species in the study plots. In the establishment year, plants transplanted in home soil produced more shoots than plants in away soil indicating that plant populations at low and high diversity developed differently over time depending on their associated soil community and/or conditions. In the second year, offspring of individuals selected at high diversity generally had a higher performance (biomass production and fitness) than offspring of individuals selected at low diversity, regardless of the transplant environment. This suggests that plants at low and high diversity showed rapid evolutionary responses measurable in their phenotype. Our findings provide first empirical evidence that loss of productivity at low diversity is not only caused by changes in abiotic and biotic conditions but also that plants respond to this by a change in their micro‐evolution. Thus, we conclude that eco‐evolutionary feedbacks of plants at low and high diversity are critical to fully understand why the positive influence of diversity on plant productivity is strengthening through time. [ABSTRACT FROM AUTHOR]

Canopy structure is an important driver of the energy budget of grassland ecosystem and is, at the same time, altered by plant diversity. Diverse plant communities typically have taller and more densely packed canopies than less diverse communities. With this, they absorb more radiation, have a higher transpiring leaf surface and are better coupled to the atmosphere which leads to cooler canopy surfaces. However, whether plant diversity generally translates into a cooling potential remains unclear and lacks empirical evidence. Here, we assessed how functional identity, functional diversity and species richness of grassland communities in the Jena Experiment predict the mean and variation of plant surface temperature mediated via effects of canopy structure.Using terrestrial laser scanning, we estimated canopy structure describing metrics of vertical structure (mean height, LAI), the distribution (evenness) and the highest allocation (centre of gravity) of biomass along height strata. As metrics of horizontal structure, we considered community stand gaps, canopy surface variation and emergent flowers. We measured surface temperature with a thermal camera. We used SEM models to predict biodiversity effects on the surface temperature during two seasonal peaks of biomass.Before the first cut in May, herb‐dominated communities directly promoted lower leaf surface temperatures. However, communities with lower centre of gravity (mostly herb dominated) also increased canopy surface temperatures compared with grass‐dominated communities with higher biomass stored in the top canopy. Grass‐dominated communities showed a smaller variation of surface temperatures, which was also positively affected by species richness via an increase in mean height. In August, mean surface temperature decreased with increasing community CC and LAI. The variation of surface temperature was greater in herb‐dominated than in grass‐dominated communities and increased with plant species richness (direct effects).Synthesis. The mean and variation of canopy surface temperature were driven by differences in functional group composition (herbs‐ vs. grass dominance) and to a lesser extent by plant diversity. These effects were partly mediated the metrics of canopy structure but also by direct effects unrelated to the structural metrics considered. [ABSTRACT FROM AUTHOR]

Although nitrogen (N) deposition is increasing globally, N availability still limits many organisms, such as microorganisms and mesofauna. However, little is known to which extent soil organisms rely on mineral‐derived N and whether plant community composition modifies its incorporation into soil food webs. More diverse plant communities more effectively compete with microorganisms for mineral N likely reducing the incorporation of mineral‐derived N into soil food webs. We set up a field experiment in experimental grasslands with different levels of plant species and functional group richness. We labeled soil with 15NH415NO3 and analyzed the incorporation of mineral‐derived 15N into soil microorganisms and mesofauna over 3 months. Mineral‐derived N incorporation decreased over time in all investigated organisms. Plant species richness and presence of legumes reduced the uptake of mineral‐derived N into microorganisms. In parallel, the incorporation of mineral‐derived 15N into mesofauna species declined with time and decreased with increasing plant species richness in the secondary decomposer springtail Ceratophysella sp. Effects of both plant species richness and functional group richness on other mesofauna species varied with time. The presence of grasses increased the 15N incorporation into Ceratophysella sp., but decreased it in the primary decomposer oribatid mite Tectocepheus velatus sarekensis. The results highlight that mineral N is quickly channeled into soil animal food webs via microorganisms irrespective of plant diversity. The amount of mineral‐derived N incorporated into soil animals, and the plant community properties affecting this incorporation, differed markedly between soil animal taxa, reflecting species‐specific use of food resources. Our results highlight that plant diversity and community composition alter the competition for N in soil and change the transfer of N across trophic levels in soil food webs, potentially leading to changes in soil animal population dynamics and community composition. Sustaining high plant diversity may buffer detrimental effects of elevated N deposition on soil biota. [ABSTRACT FROM AUTHOR]

Plant diversity is an important driver of below‐ground ecosystem functions, such as root growth, soil organic matter (SOM) storage and microbial metabolism, mainly by influencing the interactions between plant roots and soil. Dissolved organic matter (DOM), as the most mobile form of SOM, plays a crucial role for a multitude of soil processes that are central for ecosystem functioning. Thus, DOM is likely to be an important mediator of plant diversity effects on soil processes. However, the relationships between plant diversity and DOM have not been studied so far.We investigated the mechanisms underlying plant diversity effects on concentrations of DOM using continuous soil water sampling across 6 years and 62 plant communities in a long‐term grassland biodiversity experiment in Jena, Germany. Furthermore, we investigated plant diversity effects on the molecular properties of DOM in a subset of the samples.Although DOM concentrations were highly variable over the course of the year with highest concentrations in summer and autumn, we found that DOM concentrations consistently increased with plant diversity across seasons. The positive plant diversity effect on DOM concentrations was mainly mediated by increased microbial activity and newly sequestered carbon in topsoil. However, the effect of soil microbial activity on DOM concentrations differed between seasons, indicating DOM consumption in winter and spring, and DOM production in summer and autumn. Furthermore, we found increased contents of small and easily decomposable DOM molecules reaching deeper soil layers with high plant diversity.Synthesis. Our findings suggest that plant diversity enhances the continuous downward transport of DOM in multiple ways. On the one hand, higher plant diversity results in higher DOM concentrations, on the other hand, this DOM is less degraded. This study indicates, for the first time, that higher plant diversity enhances the downward transport of dissolved molecules that likely stimulate soil development in deeper layers and therefore increase soil fertility. [ABSTRACT FROM AUTHOR]

Plant diversity has a strong impact on a plethora of ecosystem functions and services, especially ecosystem carbon (C) storage. However, the potential context-dependency of biodiversity effects across ecosystem types, environmental conditions and carbon pools remains largely unknown. In this study, we performed a meta-analysis by collecting data from 95 biodiversityecosystem functioning (BEF) studies across 60 sites to explore the effects of plant diversity on different C pools, including aboveground and belowground plant biomass, soil microbial biomass C and soil C content across different ecosystem types. The results showed that ecosystem C storage was significantly enhanced by plant diversity, with stronger effects on aboveground biomass than on soil C content. Moreover, the response magnitudes of ecosystem C storage increased with the level of species richness and experimental duration across all ecosystems. The effects of plant diversity were more pronounced in grasslands than in forests. Furthermore, the effects of plant diversity on belowground plant biomass increased with aridity index in grasslands and forests, suggesting that climate change might modulate biodiversity effects, which are stronger under wetter conditions but weaker under more arid conditions. Taken together, these results provide novel insights into the important role of plant diversity in ecosystem C storage across critical C pools, ecosystem types and environmental contexts. [ABSTRACT FROM AUTHOR]

We conducted a 15N tracer experiment in laboratory microcosms with field-fresh soil samples from a biodiversity experiment to evaluate the relationship between grassland biodiversity and N cycling. To embrace the complexity of the N cycle, we determined N exchange between five soil N pools (labile and recalcitrant organic N, dissolved NH4+ and NO3− in soil solution, and exchangeable NH4+) and eight N transformations (gross N mineralization from labile and recalcitrant organic N, NH4+ immobilization into labile and recalcitrant organic N, autotrophic nitrification, heterotrophic nitrification, NO3− immobilization, adsorption of NH4+) expected in aerobic soils with the help of the N-cycle model Ntrace. We used grassland soil of the Jena Experiment, which includes plant mixtures with 1 to 60 species and 1 to 4 functional groups (legumes, grasses, tall herbs, small herbs). The 19 soil samples of one block of the Jena Experiment were labeled with either 15NH4+ or 15NO3- or both. In the presence of legumes, gross N mineralization and autotrophic nitrification increased significantly because of higher soil N concentrations in legume-containing plots and high microbial activity. Similarly, the presence of grasses significantly increased the soil NH4+ pool, gross N mineralization, and NH4+ immobilization, likely because of enhanced microbial biomass and activity by providing large amounts of rhizodeposits through their dense root systems. In our experiment, previously reported plant species richness effects on the N cycle, observed in a larger-scale field experiment within the Jena Experiment, were not seen. However, specific plant functional groups had a significant positive impact on the N cycling in the incubated soil samples. [ABSTRACT FROM AUTHOR]

Climate and land-use change lead to decreasing invertebrate biomass and alter invertebrate communities. These biotic changes may affect plant species abundance and phenology. Using 24 controlled experimental units in the iDiv Ecotron, we assessed the effects of invertebrate decline on an artificial grassland community formed by 12 herbaceous plant species. More specifically, we used Malaise traps and sweep nets to collect invertebrates from a local tall oatgrass meadow and included them in our Ecotron units at two different invertebrate densities: 100% (no invertebrate decline) and 25% (invertebrate decline of 75%). Another eight EcoUnits received no fauna and served as a control. Plant species abundance and flowering phenology was observed weekly over a period of 18 weeks. Our results showed that invertebrate densities affected the abundance and phenology of plant species. We observed a distinct species abundance shift with respect to the invertebrate treatment. Notably, this shift included a reduction in the abundance of the dominant plant species, Trifolium pratense , when invertebrates were present. Additionally, we found that the species shifted their flowering phenology as a response to the different invertebrate treatments, e.g. with decreasing invertebrate biomass Lotus corniculatus showed a later peak flowering time. We demonstrated that in addition to already well-studied abiotic drivers, biotic components may also drive phenological changes in plant communities. This study clearly suggests that invertebrate decline may contribute to already observed mismatches between plants and animals, with potential negative consequences for ecosystem services like food provision and pollination success. This deterioration of ecosystem function could enhance the loss of insects and plant biodiversity. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Aim: Nutrient enrichment is associated with plant invasions and biodiversity loss. Functional trait advantages may predict the ascendancy of invasive plants following nutrient enrichment but this is rarely tested. Here, we investigate (a) whether dominant native and non‐native plants differ in important morphological and physiological leaf traits, (b) how their traits respond to nutrient addition, and (c) whether responses are consistent across functional groups. Location: Australia, Europe, North America and South Africa. Time period: 2007–2014. Major taxa studied: Graminoids and forbs. Methods: We focused on two types of leaf traits connected to resource acquisition: morphological features relating to light‐foraging surfaces and investment in tissue (specific leaf area, SLA) and physiological features relating to internal leaf chemistry as the basis for producing and utilizing photosynthate. We measured these traits on 503 leaves from 151 dominant species across 27 grasslands on four continents. We used an identical nutrient addition treatment of nitrogen (N), phosphorus (P) and potassium (K) at all sites. Sites represented a broad range of grasslands that varied widely in climatic and edaphic conditions. Results: We found evidence that non‐native graminoids invest in leaves with higher nutrient concentrations than native graminoids, particularly at sites where native and non‐native species both dominate. We found little evidence that native and non‐native forbs differed in the measured leaf traits. These results were consistent in natural soil fertility levels and nutrient‐enriched conditions, with dominant species responding similarly to nutrient addition regardless of whether they were native or non‐native. Main conclusions: Our work identifies the inherent physiological trait advantages that can be used to predict non‐native graminoid establishment, potentially because of higher efficiency at taking up crucial nutrients into their leaves. Most importantly, these inherent advantages are already present at natural soil fertility levels and are maintained following nutrient enrichment. [ABSTRACT FROM AUTHOR]

Biodiversity is a major driver of numerous ecosystem functions. However, consequences of changes in forest biodiversity remain difficult to predict because of limited knowledge about how tree diversity influences ecosystem functions. Litter decomposition is a key process affecting nutrient cycling, productivity, and carbon storage and can be influenced by plant biodiversity. Leaf litter species composition, environmental conditions, and the detritivore community are main components of the decomposition process, but their complex interactions are poorly understood. In this study, we tested the effect of tree functional diversity (FD) on litter decomposition in a field experiment manipulating tree diversity and partitioned the effects of litter physiochemical diversity and the detritivore community. We used litterbags with different mesh sizes to separate the effects of microorganisms and microfauna, mesofauna, and macrofauna and monitored soil fauna using pitfall traps and earthworm extractions. We hypothesized that higher tree litter FD accelerates litter decomposition due to the availability of complementary food components and higher activity of detritivores. Although we did not find direct effects of tree FD on litter decomposition, we identified key litter traits and macrodetritivores that explained part of the process. Litter mass loss was found to decrease with an increase in leaf litter carbon:nitrogen ratio. Moreover, litter mass loss increased with an increasing density of epigeic earthworms, with most pronounced effects in litterbags with a smaller mesh size, indicating indirect effects. Higher litter FD and litter nutrient content were found to increase the density of surface‐dwelling macrofauna and epigeic earthworm biomass. Based on structural equation modeling, we conclude that tree FD has a weak positive effect on soil surface litter decomposition by increasing the density of epigeic earthworms and that litter nitrogen‐related traits play a central role in tree composition effects on soil fauna and decomposition. [ABSTRACT FROM AUTHOR]

The extensive use of traits in ecological studies over the last few decades to predict community functions has revealed that plant traits are plastic and respond to various environmental factors. These plant traits are assumed to predict how plants compete and capture resources. Variation in stoichiometric ratios both within and across species reflects resource capture dynamics under competition. However, the impact of local plant diversity on species‐specific stoichiometry remains poorly studied. Here, we analyze how spatial and temporal diversity in resource‐acquisition traits affects leaf elemental stoichiometry of plants (i.e. the result of resource capture) and how flexible this stoichiometry is depending on the functional composition of the surrounding community. Therefore, we assessed inter‐ and intraspecific variations of leaf carbon (C), nitrogen (N), and phosphorus (P) (and their ratios) of 20 grassland species in a large trait‐based plant diversity experiment located in Jena (Germany) by measuring leaf elemental concentrations at the species‐level along a gradient in plant trait dissimilarity. Our results show that plants showed large intra‐ and interspecific variation in leaf stoichiometry, which was only partly explained by the functional group identity (grass or herb) of the species. Elemental concentrations (N, P, but not C) decreased with plant species richness, and species tended to become more deviant from their monoculture stoichiometry with increasing trait dissimilarity in the community. These responses differed among species, some consistently increased or decreased in P and N concentrations; for other species, the negative or positive change in P and N concentrations increased with increasing trait difference between the target species and the remaining community. The strength of this relationship was significantly associated to the relative position of the species along trait gradients related to resource acquisition. Trait‐difference and trait‐diversity thus were important predictors of how species’ resource capture changed in competitive neighbourhoods. [ABSTRACT FROM AUTHOR]

Abstract: Decomposition is a vital process underlying many ecosystem functions. Although a growing number of studies have tested how litter richness affects the decomposition of above‐ground plant organs, knowledge remains limited about the decomposition of root mixtures. Here, we used a field experiment in a subtropical forest to investigate how species richness in root litter mixtures (air‐dried fresh fine roots) affects the decomposition of root litter material. On the basis of the concept of resource complementarity, we hypothesized that root litter would decompose faster as the richness of the root litter mixture increased. In addition, we expected the presence of detritivores would modify the effect of root richness on mass loss, because detritivores might experience bottom‐up effects from specific plant species and might affect microbial decomposer communities. We found that the richness level of root litter mixtures did not affect mass loss in the absence of detritivores. In the presence of detritivores, all root litter types decomposed faster. Notably, the positive effect of detritivores was stronger at low root litter richness than at high root litter richness, particularly during the early stages of decomposition (the first two sampling points) when litter mass loss was roughly double at low root litter richness compared to that at high root litter richness. The composition of the root fungal community measured at the last sampling point did not differ significantly across root richness levels, and was not affected by the presence of detritivores. Synthesis. Our findings demonstrate that detritivores modify the relationship between root litter diversity and root litter decomposition in subtropical forest ecosystems. This highlights an importance of cascade effects between different trophic organisms on ecosystem functioning. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The positive relationship between plant diversity and ecosystem functioning is likely to be co-determined by aboveground-belowground multitrophic interactions. Considering and manipulating such interactions thus is likely to significantly improve the mechanistic understanding of BEF relationships. The present proposal comprehensively investigates long-term (>4 years) plant diversity effects on soil microorganisms, nematodes, and other soil invertebrates across different ecosystems (grassland and forest ecosystems) and global change contexts (elevated [CO2 ], N deposition, warming, and drought) to identify general mechanisms. Complementary and well-directed laboratory experiments will be conducted to simulate soil feedback effects resulting from plant diversity-induced changes in soil food webs. This novel approach will allow investigating the balance between negative and positive plant-soil feedback effects and the consequences for ecosystem functioning. This holistic knowledge of changes in and interactions of above- and belowground processes is crucial to predict the long-term consequences of plant community simplification for ecosystem functioning. Experimental work will be complemented with the meta-analysis of previous work in order to reconcile prior inconsistent findings. The main objective of the present proposal is to disentangle the driving forces of plant diversity effects on soil biota as well as subsequent positive and negative feedback effects on plants. In order to achieve this, the present project has four major goals: (1) investigate long-term plant diversity effects on soil biota and functions across multiple settings in order to derive general conclusions; (2) investigate the significance of plant diversity-induced positive and negative soil feedback effects on plant performance; (3) investigate if anthropogenic stressor effects reinforce plant diversity effects on soil biota and subsequent soil feedback effects; and (4) synthesize results and perform meta-analyses to understand and reconcile inconsistent findings of previous studies on plant diversity effects on soil biota, and relate subsequent changes in soil food webs to alterations in ecosystem functioning. [ABSTRACT FROM AUTHOR]

Plant diversity effects on ecosystem functioning usually have been studied from a plant perspective. However, the mechanisms underlying biodiversity-ecosystem functioning relationships may also depend on positive or negative interactions between plants and other biotic and abiotic factors, which remain poorly understood. Here we assessed whether plant-herbivore and/or plant-detritivore interactions modify the biodiversity-ecosystem functioning relationship and the mechanisms underlying biodiversity effects, including complementarity and selection effects, biomass allocation, vertical distribution of roots, and plant survival using a microcosm experiment. We also evaluated to what extent trophic and non-trophic interactions are affected by abiotic conditions by studying drought effects. Our results show that biotic and abiotic conditions influence the shape of the biodiversity-ecosystem function relationship, varying from hump-shaped to linear. For instance, total biomass increased linearly with plant richness in the presence of detritivores, but not in the absence of detritivores. Moreover, detritivore effects on belowground plant productivity were highly context dependent, varying in the presence of herbivores. Plant interactions with soil biota, especially with herbivores, influenced the mechanisms underlying diversity effects. Herbivores increased plant complementarity and modified biomass allocation and vertical distribution of roots. Furthermore, biotic-abiotic interactions influenced plant productivity differently across plant functional groups. Our findings emphasize the importance of complex biotic interactions underlying biodiversity effects, and that these biotic interactions may change with abiotic conditions. Despite minor changes in productivity in the short-term, soil biota-induced changes in plant-plant interactions and plant survival are likely to have significant long-term consequences for ecosystem functioning. Considering the context-dependency of multichannel interactions may contribute to reconciling differences among observed patterns in biodiversity studies. Further, abiotic conditions modified the effects of biotic interactions, suggesting that changes in environmental conditions may not only affect ecosystems directly, but also change the biotic composition of and dynamics within ecosystems. [ABSTRACT FROM AUTHOR]

Niche complementarity and competitive disparity are driving mechanisms behind plant community assembly and productivity. Consequently, there is great interest in predicting species complementarity and their competitive differences from their functional traits as dissimilar species may compete less and result in more complete use of resources., Here we assessed the role of trait dissimilarities for species complementarity and competitive disparities within an experimental gradient of plant species richness and functional trait dissimilarity. Communities were assembled using three pools of grass and forb species based on a priori knowledge of traits related to (1) above- and below-ground spatial differences in resource acquisition, (2) phenological differences or (3) both. Complementarity and competitive disparities were assessed by partitioning the overyielding in mixed species communities into species complementarity and dominance effects., Community overyielding and the underlying complementarity and competitive dominance varied strongly among the three plant species pools. Overyielding and complementarity were greatest among species that were assembled based on their variation in both spatial and phenological traits. Competitive dominance was greatest when species were assembled based on spatial resource acquisition traits alone., In communities that were assembled based on species variation in only spatial or phenological traits, greater competitive dominance was predicted by greater differences in SLA and flowering initiation respectively, while greater complementarity was predicted by greater dissimilarity in leaf area and flowering senescence respectively. Greater differences in leaf area could also be linked to greater species complementarity in communities assembled based on variation in both phenological and spatial traits, but trait dissimilarity was unrelated to competitive dominance in these communities., Our results indicate that complementarity and competitive disparity among species are both driven by trait dissimilarities. However, the identity of the traits that drives the complementarity and competitive disparity depends on the trait variation among species that comprise the community. Moreover, we demonstrate that communities assembled with the greater variation in both spatial and phenological traits show the greatest complementarity among species., A is available for this article. [ABSTRACT FROM AUTHOR]

Plant diversity influences many ecosystem functions including root decomposition. However, due to the presence of multiple pathways via which plant diversity may affect root decomposition, our mechanistic understanding of their relationships is limited. In a grassland biodiversity experiment, we simultaneously assessed the effects of three pathways-root litter quality, soil biota, and soil abiotic conditions-on the relationships between plant diversity (in terms of species richness and the presence/absence of grasses and legumes) and root decomposition using structural equation modeling. Our final structural equation model explained 70% of the variation in root mass loss. However, different measures of plant diversity included in our model operated via different pathways to alter root mass loss. Plant species richness had a negative effect on root mass loss. This was partially due to increased Oribatida abundance, but was weakened by enhanced root potassium (K) concentration in more diverse mixtures. Equally, grass presence negatively affected root mass loss. This effect of grasses was mostly mediated via increased root lignin concentration and supported via increased Oribatida abundance and decreased root K concentration. In contrast, legume presence showed a net positive effect on root mass loss via decreased root lignin concentration and increased root magnesium concentration, both of which led to enhanced root mass loss. Overall, the different measures of plant diversity had contrasting effects on root decomposition. Furthermore, we found that root chemistry and soil biota but not root morphology or soil abiotic conditions mediated these effects of plant diversity on root decomposition. [ABSTRACT FROM AUTHOR]

Increasing frequency of extreme climatic events can disrupt ecosystem processes and destabilize ecosystem functioning. Biodiversity may dampen these negative effects of environmental perturbations to provide greater ecosystem stability. We assessed the effects of plant diversity on the resistance, recovery and stability of experimental grassland ecosystems in response to recurring summer drought over 7 yr. Plant biomass production was reduced during the summer drought treatment compared with control plots. However, the negative effect of drought was relatively less pronounced at high than at low plant diversity, demonstrating that biodiversity increased ecosystem resistance to environmental perturbation. Furthermore, more diverse plant communities compensated for the reduced productivity during drought by increasing spring productivity compared to control plots. The drought-induced compensatory recovery led to increased short-term variations in productivity across growing seasons in more diverse communities that stabilized the longer-term productivity across years. Our findings show that short-term variation between seasons in the face of environmental perturbation can lead to longer-term stability of annual productivity in diverse ecosystems compared to less diverse ecosystems. [ABSTRACT FROM AUTHOR]

Aims: Recent studies in experimental grasslands indicated that declining plant species diversity negatively affects soil microbial communities. Here, we assessed if plant diversity effects also occur in 'real-world' grasslands. Methods: We studied the influence of fertilization, soil, and plant community characteristics on soil microbial activity (microbial biomass carbon, basal respiration) in 12 managed temperate grasslands of varying plant species richness in two subsequent years. Results: The most important variable explaining variation in microbial activity was soil water content, while positive effects of other soil characteristics (organic carbon and nitrogen concentrations) and fertilization became more important in one study year with generally moister soil conditions. Under moister conditions, fertilization also indirectly influenced soil microbial biomass C via negative effects on plant species richness, which itself increased soil microbial biomass C. Conclusion: Our results show that variation in soil microbial activity in managed grasslands involves direct effects of fertilization as well as indirect effects through changes in plant diversity and the amount of carbon and nitrogen stored in plants and soil. These results emphasize that increased nutrient inputs in grasslands entail complex changes in ecosystem processes and indicate that mechanisms driving soil microbial activity in experimental grasslands also apply to 'real-world' grasslands. [ABSTRACT FROM AUTHOR]

Plant diversity is known to influence the abundance and diversity of belowground biota; however, patterns are not well predictable and there is still much unknown about the driving mechanisms. We analyzed changes in soil nematode community composition as affected by long-term manipulations of plant species and functional group diversity in a field experiment with plant species diversity controlled by sowing a range of 1–60 species mixtures and controlling non-sown species by hand weeding. Nematode communities contain a variety of species feeding on bacteria, fungi, plants, invertebrates, while some are omnivorous. We analyzed responses of nematode abundance and diversity to plant species and functional diversity, and used structural equation modeling (SEM) to explore the possible mechanisms underlying the observed patterns. The abundance of individuals of all nematode feeding types, except for predatory nematodes, increased with both plant species and plant functional group diversity. The abundance of microbial-feeding nematodes was related positively to aboveground plant community biomass, whereas abundance of plant-feeding nematodes was related positively to shoot C:N ratio. The abundance of predatory nematodes, in turn, was positively related to numbers of plant-feeding nematodes, but not to the abundance of microbial feeders. Interestingly, the numbers of plant-feeding nematodes per unit root mass were lowest in the high-diversity plant communities, pointing at reduced exposure to belowground herbivores when plants grow in species-diverse communities. Taxon richness of plant-feeding and microbialfeeding nematodes increased with plant species and plant functional group diversity. Increasing plant functional group diversity also enhanced taxon richness of predatory nematodes. The SEM suggests that bottom-up control effects of plant species and plant functional group diversity on abundance of nematodes in the various feeding types predominantly involve mechanistic linkages related to plant quality instead of plant quantity; especially, C:N ratios of the shoot tissues, and/or effects of plants on the soil habitat, rather than shoot quantity explained nematode abundance. Although aboveground plant properties may only partly serve as a proxy for belowground resource quality and quantity, our results encourage further studies on nematode responses to variations in plant species and plant functional diversity in relation to both quantity and quality of the belowground resources. [ABSTRACT FROM AUTHOR]

Higher plant diversity reduces nitrate leaching by complementary resource use, while its relation to leaching of other N species is unclear. We determined the effects of plant species richness, functional group richness, and the presence of specific functional groups on ammonium, dissolved organic N (DON), and total dissolved N (TDN) leaching from grassland in the first 4 years after conversion from fertilized arable land to unfertilized grassland. On 62 experimental plots in Jena, Germany, with 1-60 plant species and 1-4 functional groups (legumes, grasses, tall herbs, small herbs), nitrate, ammonium, and TDN concentrations in soil solution (0-0.3 m soil layer) were measured fortnightly during 4 years. DON concentrations were calculated by subtracting inorganic N from TDN. Nitrogen concentrations were multiplied with modeled downward water fluxes to obtain N leaching. DON leaching contributed most to TDN leaching (64 ± SD 4% of TDN). Ammonium leaching was unaffected by plant diversity. Increasing species richness decreased DON leaching in the fourth year. We attribute this finding to enhanced use of DON as a C and N source and enhanced mineralization of DON by soil microorganisms. An increase of species richness decreased TDN leaching likely driven by the complementary use of nitrate by diverse mixtures. Legumes increased DON and TDN leaching likely because of their N $$_{2}$$ -fixing ability and higher litter production. Grasses decreased TDN leaching because of more exhaustive use of nitrate and water. Our results demonstrate that increasing plant species richness decreases leaching of DON and TDN. [ABSTRACT FROM AUTHOR]

Stable provisioning of ecosystem functions and services is crucial for human well-being in a changing world. Two essential ecological components driving vital ecosystem functions in terrestrial ecosystems are plant diversity and soil microorganisms. In this study, we tracked soil microbial basal respiration and biomass over a time period of 12 years in a grassland biodiversity experiment (the Jena Experiment) and examined the role of plant diversity and plant functional group composition for the spatial and temporal stability of soil microbial properties (basal respiration and biomass) in bulk-soil. Spatial and temporal stability were calculated as the inverse coefficient of variation (CV−1) of soil microbial respiration and biomass measured from soil samples taken over space and time, respectively. We found that 1) plant species richness consistently increased soil microbial properties after a time lag of four years since the establishment of the experimental plots, 2) plant species richness had minor effects on the spatial stability of soil microbial properties, whereas 3) the functional composition of plant communities significantly affected spatial stability of soil microbial properties, with legumes and tall herbs reducing both the spatial stability of microbial respiration and biomass, while grasses increased the latter, and 4) the effect of plant diversity on temporal stability of soil microbial properties turned from being negative to neutral, suggesting that the recovery of soil microbial communities from former arable land-use takes more than a decade. Our results highlight the importance of plant functional group composition for the spatial and temporal stability of soil microbial properties, and hence for microbially-driven ecosystem processes, such as decomposition and element cycling, in temperate semi-natural grassland. [ABSTRACT FROM AUTHOR]

Flooding frequency is predicted to increase during the next decades, calling for a better understanding of impacts on terrestrial ecosystems and for developing strategies to mitigate potential damage. Plant diversity is expected to buffer flooding effects by providing a broad range of species’ responses. Here we report on the response of soil processes to a severe summer flood in 2013, which affected major parts of central Europe. We compared soil microbial respiration, biomass, nutrient limitation and enzyme activity in a grassland biodiversity experiment in Germany before flooding, one week and three months after the flood. Microbial biomass was reduced in the severely flooded plots at high, but not at low plant functional group richness. Flooding alleviated microbial nitrogen limitation, presumably due the input of nutrient-rich sediments. Further, the activity of soil enzymes including 1,4-β-N-acetylglucosaminidase, phenol oxidase and peroxidase increased with flooding severity, suggesting increased chitin and lignin degradation as a consequence of the input of detritus in sediments. Flooding effects were enhanced at higher plant diversity, indicating that plant diversity temporarily reduces stability of soil processes during flooding. The long-term impacts, however, remain unknown and deserve further investigation. [ABSTRACT FROM AUTHOR]

In a recent Forum paper, Wardle ( Journal of Vegetation Science, 2016) questions the value of biodiversity-ecosystem function ( BEF) experiments with respect to their implications for biodiversity changes in real world communities. The main criticism is that the previous focus of BEF experiments on random species assemblages within each level of diversity has 'limited the understanding of how natural communities respond to biodiversity loss.' He concludes that a broader spectrum of approaches considering both non-random gains and losses of diversity is essential to advance this field of research. Wardle's paper is timely because of recent observations of frequent local and regional biodiversity changes across ecosystems. While we appreciate that new and complementary experimental approaches are required for advancing the field, we question criticisms regarding the validity of BEF experiments. Therefore, we respond by briefly reiterating previous arguments emphasizing the reasoning behind random species composition in BEF experiments. We describe how BEF experiments have identified important mechanisms that play a role in real world ecosystems, advancing our understanding of ecosystem responses to species gains and losses. We discuss recent examples where theory derived from BEF experiments enriched our understanding of the consequences of biodiversity changes in real world ecosystems and where comprehensive analyses and integrative modelling approaches confirmed patterns found in BEF experiments. Finally, we provide some promising directions in BEF research. [ABSTRACT FROM AUTHOR]

Background: Several studies have examined the effects of plant colonization on aboveground communities and processes. However, the effects of plant colonization on soil microbial communities are less known. We addressed this gap by studying effects of plant colonization within an experimental plant diversity gradient in subplots that had not been weeded for 2 and 5 years. This study was part of a long-term grassland biodiversity experiment (Jena Experiment) with a gradient in plant species richness (1, 2, 4, 8, 16, and 60 sown species per plot). We measured plant species richness and productivity (aboveground cover and biomass) as well as soil microbial basal respiration and biomass in non-weeded subplots and compared the results with those of weeded subplots of the same plots. Results: After 2 and 5 years of plant colonization, the number of colonizing plant species decreased with increasing plant diversity, i.e., low-diversity plant communities were most vulnerable to colonization. Plant colonization offset the significant relationship between sown plant diversity and plant biomass production. In line with plant community responses, soil basal respiration and microbial biomass increased with increasing sown plant diversity in weeded subplots, but soil microbial properties converged in non-weeded subplots and were not significantly affected by the initial plant species richness gradient. Conclusion: Colonizing plant species change the quantity and quality of inputs to the soil, thereby altering soil microbial properties. Thus, plant community convergence is likely to be rapidly followed by the convergence of microbial properties in the soil. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]