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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. EEA Santa Cruz Fil: Ladouceur, Emma. German Centre for Integrative Biodiversity Research (iDiv); Alemania Fil: Ladouceur, Emma. Helmholtz Centre for Environmental Research – UFZ. Department of Physiological Diversity; Alemania Fil: Ladouceur, Emma. University of Leipzig. Department of Biology; Alemania Fil: Ladouceur, Emma. Martin Luther University Halle-Wittenberg. Institute of Computer Science; Alemania Fil: Blowes, Shane A. German Centre for Integrative Biodiversity Research (iDiv); Alemania Fil: Blowes, Shane A. Martin Luther University Halle-Wittenberg. Institute of Computer Science; Alemania Fil: Chase, Jonathan M. German Centre for Integrative Biodiversity Research (iDiv); Alemania Fil: Chase, Jonathan M. Martin Luther University Halle-Wittenberg. Institute of Computer Science; Alemania Fil: Clark, Adam T. German Centre for Integrative Biodiversity Research (iDiv); Alemania Fil: Clark, Adam T. Helmholtz Centre for Environmental Research – UFZ. Department of Physiological Diversity; Alemania Fil: Clark, Adam T. Karl-Franzens University of Graz. Institute of Biology; Austria. Fil: Garbowski, Magda. German Centre for Integrative Biodiversity Research (iDiv); Alemania Fil: Garbowski, Magda. Helmholtz Centre for Environmental Research – UFZ. Department of Physiological Diversity; Alemania Fil: Alberti, Juan. Universidad Nacional de Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Laboratorio de Ecología. Mar del Plata; Argentina. Fil: Alberti, Juan. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Fil: Arnillas, Carlos Alberto. University of Toronto. Department of Physical and Environmental Sciences; Canadá. Fil: Bakker, Jonathan D. University of Washington. School of Environmental and Forest Sciences; Estados Unidos Fil: Barrio, Isabel C. Agricultural University of Iceland. Faculty of Environmental and Forest Sciences; Islandia Fil: Bharath, Siddharth. Atria University; India. Fil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Santa Cruz; Argentina. Fil: Peri, Pablo Luis. Universidad Nacional de la Patagonia Austral; Argentina. Fil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Fil: Harpole, Stanley. German Centre for Integrative Biodiversity Research (iDiv); Alemania Fil: Harpole, Stanley. Helmholtz Centre for Environmental Research – UFZ. Department of Physiological Diversity; Alemania Martin Luther University Halle-Wittenberg. Institute of Computer Science; Alemania

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.

In many grasslands, species with specific traits occupy unique temporal positions within communities. Such intra-annual segregation is predicted to be greatest in systems with high intra-annual climate variability because fluctuating environmental conditions provide opportunities for temporal niche partitioning among species. However, because most studies on intra-annual community dynamics have been conducted at individual sites, relationships between intra-annual climate variability and seasonal community dynamics at global scales have not yet been identified. Furthermore, the same characteristics that promote species-specific responses to fluctuations in environmental conditions may also drive species-specific responses to global change drivers such as eutrophication. Research provides evidence that eutrophication alters inter-annual plant community dynamics yet understanding of how it alters intra-annual dynamics remains limited.We used early-season and late-season compositional data collected from 10 grassland sites around the world to ask how intra-annual variability in precipitation and temperature as well as nutrient enrichment shape intra-annual species segregation, or seasonal β-diversity, in plant communities. We also assessed whether changes in the abundances of specific functional groups including annual forbs, perennial forbs, C3 and C4 graminoids, and legumes underpin compositional differences between early- and late-season communities and treatments. We found that intra-annual temperature variability and seasonal β-diversity were positively related but observed no relationship between intra-annual precipitation variability and seasonal β-diversity. This suggests that positive relationships between α-diversity and intra-annual temperature variability identified in earlier studies may be underpinned by the positive influence of intra-annual temperature variability on temporal segregation of species within growing seasons. We found that nutrient enrichment increased seasonal β-diversity via increased turnover of species between early- and late-season communities. This finding mirrors patterns observed at inter-annual scales and suggests fertilization can alter compositional dynamics via similar mechanisms at varied temporal scales. Finally, fertilization reduced the abundance of C4 graminoids and legumes and eliminated intra-annual differences in these groups. In contrast, fertilization resulted in intra-annual differences in C3 graminoids which were not observed in control conditions, and increased abundance of C3 graminoids and annual forbs overall. Our study provides new insight into how intra-annual climate variability and nutrient enrichment influence biodiversity and seasonal dynamics in global grasslands.

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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.

Plant–soil feedback (PSF) has gained attention as a mechanism promoting plant growth and coexistence. However, most PSF research has measured monoculture growth in greenhouse conditions. Translating PSFs into effects on plant growth in field communities remains an important frontier for PSF research. Using a 4‐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., 72 PSFs). Plant community models were parameterized with or without these PSF effects, and model predictions were compared to plant biomass production in diversity–productivity experiments. Plants created soils that changed subsequent plant biomass by 40%. However, because they were both positive and negative, the average PSF effect was 14% less growth on “home” than on “away” soils. Nine‐species plant communities produced 29 to 37% more biomass for polycultures than for monocultures due primarily to selection effects. With or without PSF, plant community models predicted 28%–29% more biomass for polycultures than for monocultures, again due primarily to selection effects. Synthesis: Despite causing 40% changes in plant biomass, PSFs had little effect on model predictions of plant community biomass across a range of species richness. While somewhat surprising, a lack of a PSF effect was appropriate in this site because species richness effects in this study were caused by selection effects and not complementarity effects (PSFs are a complementarity mechanism). Our plant community models helped us describe several reasons that even large PSF may not affect plant productivity. Notably, we found that dominant species demonstrated small PSF, suggesting there may be selective pressure for plants to create neutral PSF. Broadly, testing PSFs in plant communities in field conditions provided a more realistic understanding of how PSFs affect plant growth in communities in the context of other species traits., Plant–soil feedbacks have gained attention as a mechanism of plant growth and coexistence, yet most plant–soil feedback research continues to be performed in greenhouse conditions. At the end of a 4‐year field experiment, plants created soils that changed subsequent plant growth by 37%, yet these relatively large effects had little effect on plant community growth because some feedbacks were positive and others negative and because feedbacks for the dominant species were small. Research highlights the need to test the consequences of plant–soil feedbacks in plant communities.

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.

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

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.

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

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.

1- Plant damage by invertebrate herbivores and pathogens influences the dynamics of grassland ecosystems, but anthropogenic changes in nitrogen and phosphorus availability can modify these relationships. 2- Using a globally distributed experiment, we describe leaf damage on 153 plant taxa from 27 grasslands worldwide, under ambient conditions and with experimentally elevated nitrogen and phosphorus. 3- Invertebrate damage significantly increased with nitrogen addition, especially in grasses and non-leguminous forbs. Pathogen damage increased with nitrogen in grasses and legumes but not forbs. Effects of phosphorus were generally weaker. Damage was higher in grasslands with more precipitation, but climatic conditions did not change effects of nutrients on leaf damage. On average, invertebrate damage was relatively higher on legumes and pathogen damage was relatively higher on grasses. Community-weighted mean damage reflected these functional group patterns, with no effects of N on community-weighted pathogen damage (due to opposing responses of grasses and forbs) but stronger effects of N on community-weighted invertebrate damage (due to consistent responses of grasses and forbs). 4- Synthesis. As human-induced inputs of nitrogen and phosphorus continue to increase, understanding their impacts on invertebrate and pathogen damage becomes increasingly important. Our results demonstrate that eutrophication frequently increases plant damage and that damage increases with precipitation across a wide array of grasslands. Invertebrate and pathogen damage in grasslands is likely to increase in the future, with potential consequences for plant, invertebrate and pathogen communities, as well as the transfer of energy and nutrients across trophic levels. EEA Santa Cruz Fil: Ebeling, Anne. University of Jena. Institute of Ecology and Evolution; Alemania Fil: Strauss, Alex T. University of Minnesota. Department of Ecology, Evolution, and Behavior; Estados Unidos Fil: Strauss, Alex T. University of Georgia. Odum School of Ecology; Estados Unidos Fil: Adler, Peter B. Utah State University. Department of Wildland Resources and the Ecology Center; Estados Unidos Fil: Arnillas, Carlos Alberto. University of Toronto —Scarborough. Department of Physical and Environmental Sciences; Canadá Fil: Barrio, Isabel C. Agricultural University of Iceland. Faculty of Environmental and Forest Sciences; Islandia Fil: Biederman, Lori A. Iowa State University. Department of Ecology, Evolution, and Organismal Biology; Estados Unidos Fil. Borer, Elizabeth T. University of Minnesota. Department of Ecology, Evolution, and Behavior; Estados Unidos Fil: Bugalho, Miguel N. University of Lisbon. Centre for Applied Ecology (CEABN-InBIO). School of Agriculture; Portugal. Fil: Caldeira, Maria C. University of Lisbon. Forest Research Centre. School of Agriculture; Portugal. Fil: Cadotte, Marc W. University of Toronto Scarborough. Department of Biological Sciences; Canadá Fil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Santa Cruz; Argentina. Fil: Peri, Pablo Luis. Universidad Nacional de la Patagonia Austral; Argentina. Fil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Fil: Blumenthal, Dana M. USDA-ARS, Rangeland Resources & Systems Research Unit; Estados Unidos

Fertilisation experiments have demonstrated that nutrient availability is a key determinant of biomass production and carbon sequestration in grasslands. However, the influence of nutrients in explaining spatial variation in grassland biomass production has rarely been assessed. Using a global dataset comprising 72 sites on six continents, we investigated which of 16 soil factors that shape nutrient availability associate most strongly with variation in grassland aboveground biomass. Climate and N deposition were also considered. Based on theory-driven structural equation modelling, we found that soil micronutrients (particularly Zn and Fe) were important predictors of biomass and, together with soil physicochemical properties and C:N, they explained more unique variation (32%) than climate and N deposition (24%). However, the association between micronutrients and biomass was absent in grasslands limited by NP. These results highlight soil properties as key predictors of global grassland biomass production and point to serial co-limitation by NP and micronutrients.

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.

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.

Droughts can strongly affect grassland productivity and biodiversity, but responses differ widely. Nutrient availability may be a critical factor explaining this variation, but is often ignored in analyses of drought responses. Here, we used a standardized nutrient addition experiment covering 10 European grasslands to test if full-factorial nitrogen, phosphorus, and potassium addition affected plant community responses to inter-annual variation in drought stress and to the extreme summer drought of 2018 in Europe. We found that nutrient addition amplified detrimental drought effects on community aboveground biomass production. Drought effects also differed between functional groups, with a negative effect on graminoid but not forb biomass production. Our results imply that eutrophication in grasslands, which promotes dominance of drought-sensitive graminoids over forbs, amplifies detrimental drought effects. In terms of climate change adaptation, agricultural management would benefit from taking into account differential drought impacts on fertilized versus unfertilized grasslands, which differ in ecosystem services they provide to society.

Earth is home to over 350,000 vascular plant species that differ in their traits in innumerable ways. A key challenge is to predict how natural or anthropogenically driven changes in the identity, abundance and diversity of co-occurring plant species drive important ecosystem-level properties such as biomass production or carbon storage. Here, we analyse the extent to which 42 different ecosystem properties can be predicted by 41 plant traits in 78 experimentally manipulated grassland plots over 10 years. Despite the unprecedented number of traits analysed, the average percentage of variation in ecosystem properties jointly explained was only moderate (32.6%) within individual years, and even much lower (12.7%) across years. Most other studies linking ecosystem properties to plant traits analysed no more than six traits and, when including only six traits in our analysis, the average percentage of variation explained in across-year levels of ecosystem properties dropped to 4.8%. Furthermore, we found on average only 12.2% overlap in significant predictors among ecosystem properties, indicating that a small set of key traits able to explain multiple ecosystem properties does not exist. Our results therefore suggest that there are specific limits to the extent to which traits per se can predict the long-term functional consequences of biodiversity change, so that data on additional drivers, such as interacting abiotic factors, may be required to improve predictions of ecosystem property levels.

Increasing plant diversity controls herbivory by limiting energetic gains and enhancing predation of arthropod herbivores., Arthropod herbivores cause substantial economic costs that drive an increasing need to develop environmentally sustainable approaches to herbivore control. Increasing plant diversity is expected to limit herbivory by altering plant-herbivore and predator-herbivore interactions, but the simultaneous influence of these interactions on herbivore impacts remains unexplored. We compiled 487 arthropod food webs in two long-running grassland biodiversity experiments in Europe and North America to investigate whether and how increasing plant diversity can reduce the impacts of herbivores on plants. We show that plants lose just under half as much energy to arthropod herbivores when in high-diversity mixtures versus monocultures and reveal that plant diversity decreases effects of herbivores on plants by simultaneously benefiting predators and reducing average herbivore food quality. These findings demonstrate that conserving plant diversity is crucial for maintaining interactions in food webs that provide natural control of herbivore pests.

Microbial processing of aggregate-unprotected organic matter inputs is key for soil fertility, long-term ecosystem carbon and nutrient sequestration and sustainable agriculture. We investigated the effects of adding multiple nutrients (nitrogen, phosphorus and potassium plus nine essential macro- and micro-nutrients) on decomposition and biochemical transformation of standard plant materials buried in 21 grasslands from four continents. Addition of multiple nutrients weakly but consistently increased decomposition and biochemical transformation of plant remains during the peak-season, concurrent with changes in microbial exoenzymatic activity. Higher mean annual precipitation and lower mean annual temperature were the main climatic drivers of higher decomposition rates, while biochemical transformation of plant remains was negatively related to temperature of the wettest quarter. Nutrients enhanced decomposition most at cool, high rainfall sites, indicating that in a warmer and drier future fertilized grassland soils will have an even more limited potential for microbial processing of plant remains. Fil: Ochoa Hueso, Raúl. Universidad de Cádiz; España Fil: Borer, Elizabeth. University of Minnesota; Estados Unidos Fil: Seabloom, Eric. University of Minnesota; Estados Unidos Fil: Hobbie, Sarah E.. University of Minnesota; Estados Unidos Fil: Risch, Anita C.. Swiss Federal Institute for Forest, Snow and Landscape Research; Suiza Fil: Collins, Scott L.. University of New Mexico; Estados Unidos Fil: Alberti, Juan. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Bahamonde, Héctor Alejandro. Universidad Nacional de la Patagonia Austral; Argentina. Instituto Nacional de Tecnología Agropecuaria; Argentina Fil: Brown, Cynthia S.. State University of Colorado - Fort Collins; Estados Unidos Fil: Caldeira, Maria C.. Universidade Nova de Lisboa; Portugal Fil: Daleo, Pedro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Dickman, Chris R.. University of Sydney; Australia Fil: Ebeling, Anne. Universitat Jena; Alemania Fil: Eisenhauer, Nico. German Centre for Integrative Biodiversity Research; Alemania. Universitat Leipzig; Alemania Fil: Esch, Ellen H.. University Of Guelph. Department Of Integrative Biology.; Canadá Fil: Eskelinen, Anu. German Centre for Integrative Biodiversity Research; Alemania. Helmholtz Centre for Environmental Research; Alemania. University of Oulu; Finlandia Fil: Fernández, Victoria. Universidad Politécnica de Madrid; España Fil: Güsewell, Sabine. Institute of Integrative Biology; Suiza Fil: Gutierrez Larruga, Blanca. Universidad Autónoma de Madrid; España Fil: Hofmockel, Kirsten. Pacific Northwest National Laboratory; Estados Unidos. University of Iowa; Estados Unidos Fil: Laungani, Ramesh. Doane University; Estados Unidos Fil: Lind, Eric. University of Minnesota; Estados Unidos Fil: López, Andrea. University of New Mexico; Estados Unidos Fil: McCulley, Rebecca L.. University of Kentucky; Estados Unidos Fil: Moore, Joslin L.. Monash University; Australia Fil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro de Investigaciones y Transferencia de Santa Cruz. Universidad Tecnológica Nacional. Facultad Regional Santa Cruz. Centro de Investigaciones y Transferencia de Santa Cruz. Universidad Nacional de la Patagonia Austral. Centro de Investigaciones y Transferencia de Santa Cruz; Argentina Fil: Power, Sally A.. University of Western Sydney; Australia Fil: Price, Jodi N.. Charles Sturt University; Australia Fil: Prober, Suzanne M.. Csiro National Collections And Marine Infrastructure; Australia Fil: Roscher, Christiane. German Centre for Integrative Biodiversity Research; Alemania. University of Oulu; Finlandia Fil: Velasco Ayuso, Sergio. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura. Universidad de Buenos Aires. Facultad de Agronomía. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura; Argentina

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.

Contains fulltext : 217065.pdf (Publisher’s version ) (Open Access) Abstract Locally, plant species richness supports many ecosystem functions. Yet, the mechanisms driving these often-positive biodiversity?ecosystem functioning relationships are not well understood. Spatial resource partitioning across vertical resource gradients is one of the main hypothesized causes for enhanced ecosystem functioning in more biodiverse grasslands. Spatial resource partitioning occurs if species differ in where they acquire resources and can happen both above- and belowground. However, studies investigating spatial resource partitioning in grasslands provide inconsistent evidence. We present the results of a meta-analysis of 21 data sets from experimental species-richness gradients in grasslands. We test the hypothesis that increasing spatial resource partitioning along vertical resource gradients enhances ecosystem functioning in diverse grassland plant communities above- and belowground. To test this hypothesis, we asked three questions. (1) Does species richness enhance biomass production or community resource uptake across sites? (2) Is there evidence of spatial resource partitioning as indicated by resource tracer uptake and biomass allocation above- and belowground? (3) Is evidence of spatial resource partitioning correlated with increased biomass production or community resource uptake? Although plant species richness enhanced community nitrogen and potassium uptake and biomass production above- and belowground, we found that plant communities did not meet our criteria for spatial resource partitioning, though they did invest in significantly more aboveground biomass in higher canopy layers in mixture relative to monoculture. Furthermore, the extent of spatial resource partitioning across studies was not positively correlated with either biomass production or community resource uptake. Our results suggest that spatial resource partitioning across vertical resource gradients alone does not offer a general explanation for enhanced ecosystem functioning in more diverse temperate grasslands.