Summary 1. Plant functional diversity and soil microbial community composition are tightly coupled. Changes in these interactions may influence ecosystem functioning. Links between plant functional diversity, soil microbial communities and ecosystem functioning have been demonstrated in experiments using plant monocultures and mixtures, using broad plant and microbial functional groups, but have not been examined in diverse natural plant communities. 2. We quantified the relative effects of plant and microbial functional properties on key ecosystem functions. We measured plant functional diversity, soil microbial community composition and parameters associated with nitrogen (N) cycling and key nutrient cycling processes at three grassland sites in different parts of Europe. Because plant structure and function strongly influence soil microbial communities, we determined relationships between ecosystem properties, plant traits and soil community characteristics following a sequential approach in which plant traits were fitted first, followed by the additional effects of soil micro-organisms. 3. We identified a continuum from standing green biomass and standing litter, linked mostly with plant traits, to potential N mineralization and potential leaching of soil inorganic N, linked mostly with microbial properties. Plant and microbial functional parameters were equally important in explaining % organic matter content in soil. A parallel continuum ran from plant height, linked with above-ground biomass, to plant quality effects captured by the leaf economics spectrum, which were linked with the recycling of carbon (C) and N. 4. More exploitative species (higher specific leaf area, leaf N concentrations and lower leaf dry matter content) and taller swards, along with soil microbial communities dominated by bacteria, with rapid microbial activities, were linked with greater fodder production, but poor C and N retention. Conversely, dominance by conservative species (with opposite traits) and soil microbial communities dominated by fungi, and bacteria with slow activities, were usually linked with low production, but greater soil C storage and N retention. 5. Synthesis -Grassland production, C sequestration and soil N retention are jointly related to plant and microbial functional traits. Managing grasslands for selected, or multiple, ecosystem services will thus require a consideration of the joint effects of plant and soil communities. Further understanding of the mechanisms that link plant and microbial functional traits is essential to achieve this.

The results suggest that consideration of plant traits, and especially below-ground traits, increases our ability to describe variation in the abundances and the functional characteristics of microbial communities in grassland soils.

Summary1. Functional variability (FV) of populations can be decomposed into three main features: the individual variability of multiple traits, the strength of correlations between those traits and the main direction of these correlations, the latter two being known as 'phenotypic integration'. Evolutionary biology has long recognized that FV in natural populations is key to determining potential evolutionary responses, but this topic has been little studied in functional ecology. 2. Here, we focus on the arctico-alpine perennial plant species Polygonum viviparum L.. We used a comprehensive sampling of seven functional traits in 29 wild populations covering the whole environmental niche of the species. The niche of the species was captured by a temperature gradient, which separated alpine stressful habitats from species-rich, competitive subalpine ones. We sought to assess the relative roles of abiotic stress and biotic interactions in shaping different aspects of functional variation within and among populations, that is, the multi-trait variability, the strength of correlations between traits and the main directions of functional trade-offs. 3. Populations with the highest extent of functional variability were found in the warm end of the gradient, whereas populations exhibiting the strongest degree of phenotypic integration were located in sites with intermediate temperatures. This could reveal both the importance of environmental filtering and population demography in structuring FV. Interestingly, we found that the main axes of multivariate functional variation were radically different within and across population. 4. Although the proximate causes of FV structure remain uncertain, our study presents a robust methodology for the quantitative study of functional variability in connection with species' niches. It also opens up new perspectives for the conceptual merging of intraspecific functional patterns with community ecology.

1. Trait-based approaches provide a framework to understand the role of functional biodiversity on ecosystem functioning under global change. While plant traits have been reported as potential drivers of soil microbial community composition and resilience, studies directly assessing microbial traits are scarce, limiting our mechanistic understanding of ecosystem functioning.2. We used microbial biomass and enzyme stoichiometry, and mass-specific enzymes activity as proxies of microbial community-weighted mean (CWM) traits, to infer trade-offs in microbial strategies of resource use with cascading effects on ecosystem resilience. We simulated a drought event on intact plant-soil mesocosms extracted from mountain grasslands along a management intensity gradient.Ecosystem processes and properties related to nitrogen cycling were quantified before, during and after drought to characterize ecosystem resilience.3. Soil microbial CWM traits and ecosystem resilience to drought were strongly influenced by grassland type. Structural equation modelling revealed a cascading effect from management to ecosystem resilience through modifications in soil nutrients, and plant and microbial CWM traits. Overall, our results depict a shift from high investment in extracellular enzymes in nutrient-poor soils (oligotrophic strategy), to a copiotrophic strategy with low microbial biomass N:P and low investment in extracellular enzymes associated with exploitative plant traits in nutrient-rich soils.4. Microbial CWM traits responses to management intensity were highly related to ecosystem resilience. Microbial communities with a copiotrophic strategy had lower resistance but higher recovery to drought, while microbial communities with an oligotrophic strategy showed the opposite responses. The unexpected trade-off between plant and microbial resistance suggested that the lower resistance of copiotrophic microbial communities enabled plant resistance to drought. Synthesis.Grassland management has cascading effects on ecosystem resilience through its combined effects on soil nutrients and plant traits propagating to microbial traits and resilience. We suggest that intensification of permanent grassland management and associated increases in soil nutrient availability decreased | 877 Journal of Ecology PITON eT al. S U PP O RTI N G I N FO R M ATI O N Additional supporting information may be found online in the Supporting Information section. How to cite this article: Piton G, Legay N, Arnoldi C, Lavorel S, Clément J-C, Foulquier A. Using proxies of microbial community-weighted means traits to explain the cascading effect of management intensity, soil and plant traits on ecosystem resilience in mountain grasslands.

International audienceBackground and aims: drought events, agricultural practices and plant communities influence microbial and soil abiotic parameters which can feedback to fodder production. This study aimed to determine which soil legacies influence plant biomass production and nutritional quality, and its resistance and recovery to extreme weather events.Methods: In a greenhouse experiment, soil legacy effects on Lolium perenne were examined, first under optimal conditions, and subsequently during and after drought. We used subalpine grassland soils previously cultivated for two years with grass communities of distinct functional composition, and subjected to combinations of climatic stress and simulated management.Results: The soil legacy of climatic stress increased biomass production of Lolium perenne and its resistance and recovery to a new drought. This beneficial effect resulted from higher nutrient availability in soils previously exposed to climatic stresses due to lower competitive abilities and resistance of microbial communities to a new drought. This negative effect on microbial communities was strongest in soils from previously cut and fertilized grasslands or dominated by conservative grasses.Conclusion: In subalpine grasslands more frequent climatic stresses could benefit fodder production in the short term, but threaten ecosystem functioning and the maintenance of traditional agricultural practices in the long term

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With the increased occurrence of climate extremes, there is an urgent need to better understand how management strategies affect the capacity of the soil microbial community to maintain its ecosystem functions (e.g. nutrient cycling). To address this issue, intact monoliths were extracted from conventional and ecological managed grasslands in three countries across Europe and exposed under common air condition (temperature and moisture) to one of three altered rain regimes (dry, wet and intermittent wet/dry) as compared to a normal regime. Subsequently, we compared the resistance and recovery of the soil microbial biomass, potential enzyme activities and community composition. The microbial community composition differed with soil management and rain regimes. Soil microbial biomass increased from the wetter to the dryer rain regime, paralleling an increase of available carbon and nutrients, suggesting low sensitivity to soil moisture reduction but nutritional limitations of soil microbes. Conversely, enzyme activities decreased with all altered rain regimes. Resistance and recovery (considering absolute distance between normal and altered rain regime) of the microbial communities depended on soil management. Conventional‐intensive management showed higher resistance of two fundamental properties for nutrient cycling (i.e. bacterial biomass and extracellular enzyme activities) yet associated with more important changes in microbial community composition. This suggests an internal community reorganization promoting biomass and activity resistance. Conversely, under ecological management bacterial biomass and enzyme activities showed better recovery capacity, whereas no or very low recovery of these properties was observed under conventional management. These management effects were consistent across the three altered rain regimes investigated, indicating common factors controlling microbial communities’ response to different climate‐related stresses. Synthesis and applications. Our study provides experimental evidence for an important trade‐off for agroecosystem management between (a) stabilizing nutrient cycling potential during an altered rain regime period at the expense of very low recovery capacity and potential long‐term effect (conventional sites) and (b) promoting the capacity of the microbial community to recover its functional potential after the end of the stress (ecological sites). Thus, management based on ecologically sound principles may be the best option to sustain long‐term soil functioning under climate change.

Ecological intensification promotes the better use of ecosystem functioning for agricultural production and as a provider of additional regulation and cultural services. We investigated the mechanisms underpinning potential ecological intensification of livestock production in the Vercors mountains (France). We quantified the variations in seven ecosystem properties associated with key ecosystem services: above-ground biomass production at first harvest, fodder digestibility, plant species richness, soil organic matter content, soil carbon content, total microbial biomass and soil bacteria:fungi ratio across 39 grassland plots representing varying management types and intensity. Our analyses confirmed joint effects of management, traits and soil abiotic parameters on variations in ecosystem properties, with the combination of management and traits being most influential. The variations explained by traits were consistent with the leaf economics spectrum model and its implications for ecosystem functioning. The observed independence between ecosystem properties relevant to production (forage biomass, digestibility and nutrient turnover) on the one hand and soil stocks (organic matter, carbon and microbial stocks) on the other hand suggests that an intensification of fodder production might be compatible with the preservation of the soil capital. We highlight that appropriate choices regarding various practices, such as the first date of grazing or mowing being dependent on soil moisture, have important consequences on a number of ecosystem properties relevant for ecosystem services and may influence biodiversity patterns. Such avenues for ecological intensification should be considered as part of further landscape- and farm-scale analyses of the relationships between farm functioning and ecosystem services.