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1. Carbon storage in vegetation and soil underpins climate regulation through carbon sequestration. Because plant species differ in their ability to capture, store and release carbon, the collective functional characteristics of plant communities (functional diversity) should be a major driver of carbon accumulation in terrestrial ecosystems. 2. Three major components of plant functional diversity could be put forward as drivers of carbon storage in ecosystems: the most abundant functional trait values, the variety of functional trait values and the abundance of particular species that could have additional effects not incorporated in the first two components. 3. We tested for associations between these components and carbon storage across 16 sites in the Chaco forest of Argentina under the same climate and on highly similar parental material. The sites differed in their plant functional diversity caused by different long-term land-use regimes. 4. We measured six plant functional traits in 27 species and weighted them by the species abundance at each site to calculate the community-weighted mean (CWM) and the functional divergence (FDvar) of each single trait and of multiple traits (FDiv). We also measured plant and soil carbon storage. Using a stepwise multiple regression analysis, we assessed which of the functional diversity components best explained carbon storage. 5. Both CWM and FDvar of plant height and wood-specific gravity, but no leaf traits, were retained as predictors of carbon storage in multiple models. Relationships of FDvar of stem traits and FDiv with carbon storage were all negative. The abundance of five species improved the predictive power of some of the carbon storage models. 6. Synthesis. All three major components of plant functional diversity contributed to explain carbon storage. What matters the most to carbon storage in these ecosystems is the relative abundance of plants with tall, and to a lesser extent dense, stems with a narrow range of variation around these values. No consistent link was found between carbon storage and the leaf traits usually associated with plant resource use strategy. The negative association of trait divergence with carbon storage provided no evidence in support to niche complementarity promoting carbon storage in these forest ecosystems.

1. Functional traits can be used to describe the composition of communities through indices that seek to explain the factors that drive community assembly, biotic effects on ecosystem processes or both. Appropriately representing functional composition is therefore essential for predicting the consequences of environmental context and management actions for the provisioning of multiple ecosystem services (ESs) in heterogeneous landscapes. 2. Functional indices can be constructed from single or multiple traits; however, it is not clear how they differ in information content or ability to predict biodiversity — ecosystem function relationships in complex landscapes. Here, we compare the utility of analogous single- and multi-trait indices in linking environmental variation and functional composition to ESs in a heterogeneous landscape, relating functional indices based on three plant traits [height, relative growth rate and root density (RD)] to variation in the physical environment and to two ESs (forage production and soil carbon) and their net ES level. 3. Two orthogonal gradients, elevation and soil bulk density (BD), explained significant variation in several dimensions of functional composition comprised of single traits. These traits in turn significantly predicted variation in ESs and their net values. Only one index measured with multiple traits (functional richness) varied with the physical environment, while none predicted variation in ES or net ES levels. 4. One ES, soil carbon, increased with the community-average value of RD, while the other, forage production, was related to the range and community-average value of height. In turn, average RD increased with soil BD while the average and range of height declined with elevation. Due to these environmental patterns, soil carbon and forage production did not covary strongly, leading to moderate net ES levels across the landscape. 5. Synthesis: Single-trait indices of functional composition best linked variation in environmental gradients with productivity and soil carbon. Because the environment—trait functioning relationships were independent of one another, the ESs were independently distributed across the landscape, providing little evidence of synergies or trade-offs. Single- and multi-trait indices contained unique information about functional composition of these communities, and both are likely to have a place in predicting variation in ESs under different scenarios.

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.

1. Plant species can influence soil biota, which in turn can influence the relative performance of plant species. These plant-soil feedbacks (PSFs) have been hypothesized to affect many community-level dynamics including species coexistence, dominance and invasion. 2. The importance of PSFs in exotic species invasion, although widely hypothesized, has been difficult to determine because invader establishment necessarily precedes invader-mediated PSFs. Here, we combine a spatial simulation model of invasion that incorporates PSFs with a meta-analysis that synthesizes published case studies describing feedbacks between pairs of native and exotic species. 3. While our spatial model confirmed the link between positive soil feedbacks ('home' advantage) for exotic species and exotic species spread, results were dependent on the initial abundance of the exotic species and the equivalence of dispersal and life history characteristics between exotic and native species. 4. The meta-analysis of 52 native-exotic pairwise feedback comparisons in 22 studies synthesized measures of native and exotic performance in soils conditioned by native and exotic species. The analysis indicated that the growth responses of native species were often greater in soil conditioned by native species than in soil conditioned exotic species (a 'home' advantage). The growth responses of exotic species were variable and not consistently related to species soil-conditioning effects. 5. Synthesis. Overlaying empirical estimates of pairwise PSFs with spatial simulations, we conclude that the empirically measured PSFs between native and exotic plant species are often not consistent with predictions of the spread of exotic species and mono-dominance. This is particularly the case when exotic species are initially rare and share similar dispersal and average fitness characteristics with native species. However, disturbance and other processes that increase the abundance of exotic species as well as the inclusion of species dispersal and life history differences can interact with PSF effects to explain the spread of invasive species.

1. Studying the effects of climate or weather extremes such as drought and heat waves on biodiversity and ecosystem functions is one of the most important facets of climate change research. In particular, primary production is amounting to the common currency in field experiments world-wide. Rarely, however, are multiple ecosystem functions measured in a single study in order to address general patterns across different categories of responses and to analyse effects of climate extremes on various ecosystem functions. 2. We set up a long-term field experiment, where we applied recurrent severe drought events annually for five consecutive years to constructed grassland communities in central Europe. The 32 response parameters studied were closely related to ecosystem functions such as primary production, nutrient cycling, carbon fixation, water regulation and community stability. 3. Surprisingly, in the face of severe drought, above- and below-ground primary production of plants remained stable across all years of the drought manipulation. 4. Yet, severe drought significantly reduced below-ground performance of microbes in soil indicated by reduced soil respiration, microbial biomass and cellulose decomposition rates as well as mycorrhization rates. Furthermore, drought reduced leaf water potential, leaf gas exchange and leaf protein content, while increasing maximum uptake capacity, leaf carbon isotope signature and leaf carbohydrate content. With regard to community stability, drought induced complementary plant-plant interactions and shifts in flower phenology, and decreased invasibility of plant communities and primary consumer abundance. 5. Synthesis. Our results provide the first field-based experimental evidence that climate extremes initiate plant physiological processes, which may serve to regulate ecosystem productivity. A potential reason for different dynamics in various ecosystem services facing extreme climatic events may lie in the temporal hierarchy of patterns of fast versus slow response. Such data on multiple response parameters within climate change experiments foster the understanding of mechanisms of resilience, of synergisms or decoupling of biogeochemical processes, and of fundamental response dynamics to drought at the ecosystem level including potential tipping points and thresholds of regime shift. Future work is needed to elucidate the role of biodiversity and of biotic interactions in modulating ecosystem response to climate extremes.

1. There is a growing consensus that the distribution of species trait values in a community can greatly determine ecosystem processes and services delivery. Two distinct components of community trait composition are hypothesized to chiefly affect ecosystem processes: (i) the average trait value of the species, quantified by community-weighted mean trait values (CWM; related to the mass ratio hypothesis) and (ii) the degree to which trait values differ between species in a community, quantified by different indices of functional diversity (FD; related to non-additive community effects). The uncertainty on the relative effect of these two components is stimulating an increasing number of empirical studies testing their effects on ecosystem processes and services delivery. 2. We suggest, however, that the interdependence between CWM and FD poses a challenge on disentangling their relative importance. We present a framework that allows designing experiments to decouple and assess the effects of these two community functional components on ecosystem processes and services. To illustrate the framework, we focused on leaf litter decomposition, as this is an essential process related to important ecosystem services. Using simulations, we applied the framework for plant leaf litter traits (litter nitrogen and phenolic content) that are related to litter decomposition. 3. CWM and FD generally showed a hump-shaped relationship (i.e. at more extreme CWM values, communities can have only low FD values). Within this relationship, we showed that it is possible to select quasi-orthogonal combinations of CWM and FD that can be treated statistically. Within these orthogonal CWM and FD combinations, it is also possible to select species assemblages controlling for other community parameters, such as total biomass, total density and species richness. 4. Synthesis. The framework provides a novel approach for designing experiments to decouple the effects of CWM and FD of communities on ecosystem processes, which otherwise cannot be easily disentangled. To apply the framework and design proper experimental layouts, it is essential to have a priori knowledge of the key traits by which species affect ecosystem processes and service delivery.

1. Recent studies have shown that the positive relationship between plant diversity and plant biomass ('over-yielding') can be explained by soil pathogens depressing productivity more in low than in high diverse plant communities. However, tests of such soil effects in field studies were constrained by experimental limitations to manipulate soil community composition independent of plant community composition. Here, we report of an experiment where feedback effects to plants were tested for both plant and soil monocultures and mixtures. 2. Our results demonstrate that overyielding is the result of plant species in mixture being more growth-limited by 'own' soil biota than by soil biota of other plant species. This effect disappeared when the soils had been sterilized by gamma-irradiation. Mixing plants themselves did not result in overyielding except when grown in the soil of one of the species (Leucanthemum vulgare), where growth of one species disproportionally increased in mixture compared to monoculture. 3. Soil nutrient availability could not explain differences in growth between the non-sterilized soils. Therefore, our results suggest that plant species-specific soil biota rather than the plants have contributed to the plant community overyielding. 4. Species biomass ranking in mixtures highly differed between non-sterilized soils of different histories of soil conditioning, whilst the ranking was more consistent in sterilized soil. Sterilized soils of different origin differed significantly in nutrient availability. These results suggest that shifts in competitive hierarchies depend on plant species-specific interactions influenced by soil biota and cannot be induced by mineral nitrogen. 5. Synthesis. Our results show that overyielding in four plant species mixtures can be due to species-specific interactions between plants and their specific soil biota. Neither mixing the plant species alone nor the differential responses of species to mineral nitrogen influenced community productivity, but mixing soil biota did.

1. Induced changes in plant quality can mediate indirect interactions between herbivores. Although the sequence of attack by different herbivores has been shown to influence plant responses, little is known about how this affects the herbivores themselves. 2. We therefore investigated how induction by the leaf herbivore Spodoptera frugiperda influences resistance of teosinte (Zea mays mexicana) and cultivated maize (Zea mays mays) against root‐feeding larvae of Diabrotica virgifera virgifera. The importance of the sequence of arrival was tested in the field and laboratory. 3. Spodoptera frugiperda infestation had a significant negative effect on colonization by D. virgifera larvae in the field and weight gain in the laboratory, but only when S. frugiperda arrived on the plant before the root herbivore. When S. frugiperda arrived after the root herbivore had established, no negative effects on larval performance were detected. Yet, adult emergence of D. virgifera was reduced even when the root feeder had established first, indicating that the negative effects were not entirely absent in this treatment. 4. The defoliation of the plants was not a decisive factor for the negative effects on root herbivore development, as both minor and major leaf damage resulted in an increase in root resistance and the extent of biomass removal was not correlated with root‐herbivore growth. We propose that leaf‐herbivore‐induced increases in feeding‐deterrent and/or toxic secondary metabolites may account for the sequence‐specific reduction in root‐herbivore performance. 5. Synthesis. Our results demonstrate that the sequence of arrival can be an important determinant of plant‐mediated interactions between insect herbivores in both wild and cultivated plants. Arriving early on a plant may be an important strategy of insects to avoid competition with other herbivores. To fully understand plant‐mediated interactions between insect herbivores, the sequence of arrival should be taken into account.

1. Indirect plant defences are well documented for the above‐ground constituents of plants. Although less investigated to date, below‐ground defences that mediate multitrophic interactions are equally important. Entomopathogenic nematodes (Steinernema diaprepesi) are attracted to herbivore‐induced volatiles from Swingle var. (Citrus paradisi×Poncirus trifoliata) when fed upon by root weevil, Diaprepes abbreviatus. 2. We examined the extent to which below‐ground volatiles modify behaviour of nematode species representing various foraging strategies (cruisers versus ambushers) and trophic levels (plant parasites versus entomopathogens). We compared attraction to volatiles of weevil‐infested and non‐infested roots from Swingle citrus rootstock and a parent line of the Swingle hybrid, Poncirus trifoliata (Pt). 3. Swingle weevil‐infested roots attracted more nematodes than non‐infested roots irrespective of nematode foraging strategy and trophic status. The parental line, Pt, attracted all nematode species irrespective of insect herbivory. 4. Dynamic in situ collection and GC-MS analysis of volatiles from soil revealed that Pt roots release attracting cues constitutively. A different non‐hybrid citrus species (sour orange, Citrus aurantium) released nematode attracting cues only in response to larval feeding, similar to responses found in Swingle. Volatile collections from above‐ and below‐ground portions of citrus plants revealed that above‐ground feeding by weevils does not induce production of nematode attracting cues analogous to that induced by root damage, nor does damage by larvae below‐ground induce a similar volatile above ground. 5. Synthesis. Our results suggest that release of nematode attractants by citrus roots occurs broadly and can be constant or herbivore‐induced. The major constituent of this indirect defence is produced by roots and not shoots and in response to below‐ground, but not above‐ground herbivory. Our findings suggest that this cue acts on nematode species broadly, attracting entomopathogenic nematodes that exhibit various foraging strategies. Unexpectedly, we also found that this cue attracts a plant parasitic nematode species. It appears, thus, that release of nematode attracting cues by citrus plants can cause ecological costs. The plants, however, appear to counteract against these costs, because constitutive release was found only in a cultivar that is resistant to phytopathogenic nematodes, while herbivore‐induced release occurred in lines susceptible to pathogenic nematode species.

1. Upon facing biotic stresses, plants orchestrate defence mechanisms via internal and external mechanisms that are mediated by signalling molecules such as salicylic acid, jasmonic acid, ethylene and various other volatile compounds. Although pathogen‐ and chemical‐induced plant resistance has been studied extensively within the same plant compartment, the effects of above‐ground (AG) insect‐elicited plant defence on the resistance expression in roots and the below‐ground (BG) microbial community are not well understood. 2. We assessed the effect of AG whitefly (Bemisia tabaci) attack on the elicitation of induced resistance against a leaf pathogen, Xanthomonas axonopodis pv. vesicatoria, a soil‐borne pathogen, Ralstonia solanacearum, and on BG modifications of the rhizosphere microflora in peppers (Capsicum annuum). 3. Symptom development caused by the two bacterial pathogens on leaves and roots was significantly reduced in whitefly‐exposed plants as compared to controls. A combined treatment with benzothiadiazole (BTH) and whitefly caused an additive effect on induced resistance, indicating that whitefly‐induced plant defence can utilize salicylic acid (SA)‐dependent signalling. To obtain further genetic evidence of this phenomenon, we evaluated the gene expression of Capsicum annuum pathogenesis‐related protein (CaPR) 1, CaPR4, CaPR10 and Ca protease inhibitor II, and observed increased expression after BTH and/or whitefly treatment indicating that AG whitefly infestation elicited SA and jasmonic acid signalling in AG and BG. Since the expression pattern of PR genes in the roots differed, we assessed microbial diversity in plants treated with BTH and/or whitefly. 4. In addition to eliciting BG defence responses, a whitefly infestation of the leaves augmented the population of root‐associated Gram‐positive bacteria and fungi, which may have positively affected plant growth and induced systemic resistance. Whitefly feeding reduced plant size, which usually occurs as a consequence of the high costs of direct resistance induction. 5. Synthesis. Our results demonstrate that whitefly‐induced resistance against bacterial pathogens can cross the AG-BG border and may cause further indirect benefits on future plant development, because it can positively affect the association or plant roots with putatively beneficial microorganisms.