Functional rarity and evenness are key facets of biodiversity to boost multifunctionality.

The functional traits of organisms within multispecies assemblages regulate biodiversity effects on ecosystem functioning. Yet how traits should assemble to boost multiple ecosystem functions simultaneously (multifunctionality) remains poorly explored. In a multibiome litter experiment covering most of the global variation in leaf trait spectra, we showed that three dimensions of functional diversity (dispersion, rarity, and evenness) explained up to 66% of variations in multifunctionality, although the dominant species and their traits remained an important predictor. While high dispersion impeded multifunctionality, increasing the evenness among functionally dissimilar species was a key dimension to promote higher multifunctionality and to reduce the abundance of plant pathogens. Because too-dissimilar species could have negative effects on ecosystems, our results highlight the need for not only diverse but also functionally even assemblages to promote multifunctionality. The effect of functionally rare species strongly shifted from positive to negative depending on their trait differences with the dominant species. Simultaneously managing the dispersion, evenness, and rarity in multispecies assemblages could be used to design assemblages aimed at maximizing multifunctionality independently of the biome, the identity of dominant species, or the range of trait values considered. Functional evenness and rarity offer promise to improve the management of terrestrial ecosystems and to limit plant disease risks.

Land-use history impacts functional diversity across multiple trophic groups.

Land-use change is a major driver of biodiversity loss worldwide. Although biodiversity often shows a delayed response to land-use change, previous studies have typically focused on a narrow range of current landscape factors and have largely ignored the role of land-use history in shaping plant and animal communities and their functional characteristics. Here, we used a unique database of 220,000 land-use records to investigate how 20-y of land-use changes have affected functional diversity across multiple trophic groups (primary producers, mutualists, herbivores, invertebrate predators, and vertebrate predators) in 75 grassland fields with a broad range of land-use histories. The effects of land-use history on multitrophic trait diversity were as strong as other drivers known to impact biodiversity, e.g., grassland management and current landscape composition. The diversity of animal mobility and resource-acquisition traits was lower in landscapes where much of the land had been historically converted from grassland to crop. In contrast, functional biodiversity was higher in landscapes containing old permanent grasslands, most likely because they offer a stable and high-quality habitat refuge for species with low mobility and specialized feeding niches. Our study shows that grassland-to-crop conversion has long-lasting impacts on the functional biodiversity of agricultural ecosystems. Accordingly, land-use legacy effects must be considered in conservation programs aiming to protect agricultural biodiversity. In particular, the retention of permanent grassland sanctuaries within intensive landscapes may offset ecological debts.

Phylogenetic, functional, and taxonomic richness have both positive and negative effects on ecosystem multifunctionality.

Biodiversity encompasses multiple attributes such as the richness and abundance of species (taxonomic diversity), the presence of different evolutionary lineages (phylogenetic diversity), and the variety of growth forms and resource use strategies (functional diversity). These biodiversity attributes do not necessarily relate to each other and may have contrasting effects on ecosystem functioning. However, how they simultaneously influence the provision of multiple ecosystem functions related to carbon, nitrogen, and phosphorus cycling (multifunctionality) remains unknown. We evaluated the effects of the taxonomic, phylogenetic, and functional attributes of dominant (mass ratio effects) and subordinate (richness effect) plant species on the multifunctionality of 123 drylands from six continents. Our results highlight the importance of the phylogenetic and functional attributes of subordinate species as key drivers of multifunctionality. In addition to a higher taxonomic richness, we found that simultaneously increasing the richness of early diverging lineages and the functional redundancy between species increased multifunctionality. In contrast, the richness of most recent evolutionary lineages and the functional and phylogenetic attributes of dominant plant species (mass ratio effects) were weakly correlated with multifunctionality. However, they were important drivers of individual nutrient cycles. By identifying which biodiversity attributes contribute the most to multifunctionality, our results can guide restoration efforts aiming to maximize either multifunctionality or particular nutrient cycles, a critical step to combat dryland desertification worldwide.

Unveiling ecological assembly rules from commonalities in trait distributions.

Deciphering the effect of neutral and deterministic processes on community assembly is critical to understand and predict diversity patterns. The information held in community trait distributions is commonly assumed as a signature of these processes, but empirical and modelling attempts have most often failed to untangle their confounding, sometimes opposing, impacts. Here, we simulated the assembly of trait distributions through stochastic (dispersal limitation) and/or deterministic scenarios (environmental filtering and niche differentiation). We characterized the shape of trait distributions using the skewness-kurtosis relationship. We identified commonalities in the co-variation between the skewness and the kurtosis of trait distributions with a unique signature for each simulated assembly scenario. Our findings were robust to variation in the composition of regional species pools, dispersal limitation and environmental conditions. While ecological communities can exhibit a high degree of idiosyncrasy, identification of commonalities across multiple communities can help to unveil ecological assembly rules in real-world ecosystems.

Biogeography of global drylands.

Despite their extent and socio-ecological importance, a comprehensive biogeographical synthesis of drylands is lacking. Here we synthesize the biogeography of key organisms (vascular and nonvascular vegetation and soil microorganisms), attributes (functional traits, spatial patterns, plant-plant and plant-soil interactions) and processes (productivity and land cover) across global drylands. These areas have a long evolutionary history, are centers of diversification for many plant lineages and include important plant diversity hotspots. This diversity captures a strikingly high portion of the variation in leaf functional diversity observed globally. Part of this functional diversity is associated with the large variation in response and effect traits in the shrubs encroaching dryland grasslands. Aridity and its interplay with the traits of interacting plant species largely shape biogeographical patterns in plant-plant and plant-soil interactions, and in plant spatial patterns. Aridity also drives the composition of biocrust communities and vegetation productivity, which shows large geographical variation. We finish our review by discussing major research gaps, which include: studying regular vegetation spatial patterns; establishing large-scale plant and biocrust field surveys assessing individual-level trait measurements; knowing whether the impacts of plant-plant and plant-soil interactions on biodiversity are predictable; and assessing how elevated CO2 modulates future aridity conditions and plant productivity.

Divergent above- and below-ground biodiversity pathways mediate disturbance impacts on temperate forest multifunctionality.

Biodiversity plays a fundamental role in provisioning and regulating forest ecosystem functions and services. Above-ground (plants) and below-ground (soil microbes) biodiversity could have asynchronous change paces to human-driven land-use impacts. Yet, we know very little how they affect the provision of multiple forest functions related to carbon accumulation, water retention capacity and nutrient cycling simultaneously (i.e. ecosystem multifunctionality; EMF). We used a dataset of 22,000 temperate forest trees from 260 plots within 11 permanent forest sites in Northeastern China, which are recovering from three post-logging disturbances. We assessed the direct and mediating effects of multiple attributes of plant biodiversity (taxonomic, phylogenetic, functional and stand structure) and soil biodiversity (bacteria and fungi) on EMF under the three disturbance levels. We found the highest EMF in highly disturbed rather than undisturbed mature forests. Plant taxonomic, phylogenetic, functional and stand structural diversity had both positive and negative effects on EMF, depending on how the EMF index was quantified, whereas soil microbial diversity exhibited a consistent positive impact. Biodiversity indices explained on average 45% (26%-58%) of the variation in EMF, whereas climate and disturbance together explained on average 7% (0.4%-15%). Our result highlighted that the tremendous effect of biodiversity on EMF, largely overpassing those of both climate and disturbance. While above- (ß = 0.02-0.19) and below-ground (ß = 0.16-0.26) biodiversity had direct positive effects on EMF, their opposite mediating effects (ß = -0.22 vs. ß = 0.35 respectively) played as divergent pathways to human disturbance impacts on EMF. Our study sheds light on the need for integrative frameworks simultaneously considering above- and below-ground attributes to grasp the global picture of biodiversity effects on ecosystem functioning and services. Suitable management interventions could maintain both plant and soil microbial biodiversity, and thus guarantee a long-term functioning and provisioning of ecosystem services in an increasing disturbance frequency world.

SGH: stress or strain gradient hypothesis? Insights from an elevation gradient on the roof of the world.

Background and Aims: The stress gradient hypothesis (SGH), the view that competition prevails in undisturbed and productive environments, and shifts to facilitation in disturbed or stressful environments, has become a central paradigm in ecology. However, an alternative view proposes that the relationship between biotic interactions and environmental severity should be unimodal instead of monotonic. Possible causes of discrepancies between these two views were examined in the high elevation desert of the arid Trans-Himalayas. Methods: A putative nurse species and its associated plant community was surveyed over its entire elevation range, spanning from alpine to desert vegetation belts. The results were analysed at the community level (vegetation cover and species richness), considering the distinction between the intensity and the importance of biotic interactions. Interactions at the species level (pairwise interactions) were also considered, i.e. the variation of biotic interactions within the niche of a species, for which the abundance (species cover) and probability of occurrence (presence/absence) for the most widespread species along the gradient were distinguished. Key Results: Overall, facilitation was infrequent in our study system; however, it was observed for the two most widespread species. At the community level, the intensity and importance of biotic interactions showed a unimodal pattern. The departure from the prediction of the SGH happened abruptly where the nurse species entered the desert vegetation belt at the lowest elevation. This abrupt shift was attributed to the turnover of species with contrasting tolerances. At the species level, however, facilitation increased consistently as the level of stress increases and individuals deviate from their optimum (increasing strain). Conclusion: While the stress gradient hypothesis was not supported along our elevation gradient at the community level, the strain gradient hypothesis, considering how species perceive the ambient level of stress and deviate from their optimum, provided a parsimonious explanation for the outcome of plant-plant interactions at both scales.

The relative contribution of short-term versus long-term effects in shrub-understory species interactions under arid conditions.

Plant-plant interactions (competition and facilitation) in terrestrial ecosystems include: (1) short-term effects, primarily quantified with experimental removals; and (2) long-term effects, mostly due to soil weathering processes, primarily quantified with observational methods. It has been argued that these effects are likely to vary in contrasting directions with increasing drought stress in arid systems. However, few studies have used appropriate methodology to assess both types of effects and their variation across nurse species and environmental conditions, in particular in arid systems. This knowledge is crucial for predicting variation in the mediating role of facilitation with climate change and assessing the importance of nurse effects in ecological restoration. In the arid climate of central-south Tunisia, understory species' biomass, abundance and composition and soil parameters were compared in shrub-control, shrub-removed and open areas for three shrub species and in two habitats with contrasting soil moisture conditions. Long-term effects were dominant, positive for all three shrub species and associated with increasing nutrient content in shrub patches. Short-term effects, mainly related to water consumption, were weaker, mostly negative and dependent on shrub species. Additionally, long-term effects were less positive and short-term effects more negative in the dry habitat than in the wet habitat. Our study provides evidence of the primary influence of positive (facilitative) long-term effects in this arid system. However, the net effects of shrubs could be less beneficial for other species with increasing aridity under climate change, due to both a decrease in positive long-term effects and an increase in negative short-term effects.

Effects of long- and short-term management on the functional structure of meadows through species turnover and intraspecific trait variability.

The functional structures of communities respond to environmental changes by both species replacement (turnover) and within-species variation (intraspecific trait variability; ITV). Evidence is lacking on the relative importance of these two components, particularly in response to both short- and long-term environmental disturbance. We hypothesized that such short- and long-term perturbations would induce changes in community functional structure primarily via ITV and turnover, respectively. To test this we applied an experimental design across long-term mown and abandoned meadows, with each plot containing a further level of short-term management treatments: mowing, grazing and abandonment. Within each plot, species composition and trait values [height, shoot biomass, and specific leaf area (SLA)] were recorded on up to five individuals per species. Positive covariations between the contribution of species turnover and ITV occurred for height and shoot biomass in response to both short- and long-term management, indicating that species turnover and intraspecific adjustments selected for similar trait values. Positive covariations also occurred for SLA, but only in response to long-term management. The contributions of turnover and ITV changed depending on both the trait and management trajectory. As expected, communities responded to short-term disturbances mostly through changes in intraspecific trait variability, particularly for height and biomass. Interestingly, for SLA they responded to long-term disturbances by both species turnover and intraspecific adjustments. These findings highlight the importance of both ITV and species turnover in adjusting grassland functional trait response to environmental perturbation, and show that the response is trait specific and affected by disturbance regime history.

Functional diversity enhances the resistance of ecosystem multifunctionality to aridity in Mediterranean drylands.

We used a functional trait-based approach to assess the impacts of aridity and shrub encroachment on the functional structure of Mediterranean dryland communities (functional diversity (FD) and community-weighted mean trait values (CWM)), and to evaluate how these functional attributes ultimately affect multifunctionality (i.e. the provision of several ecosystem functions simultaneously). Shrub encroachment (the increase in the abundance/cover of shrubs) is a major land cover change that is taking place in grasslands worldwide. Studies conducted on drylands have reported positive or negative impacts of shrub encroachment depending on the functions and the traits of the sprouting or nonsprouting shrub species considered. FD and CWM were equally important as drivers of multifunctionality responses to both aridity and shrub encroachment. Size traits (e.g. vegetative height or lateral spread) and leaf traits (e.g. specific leaf area and leaf dry matter content) captured the effect of shrub encroachment on multifunctionality with a relative high accuracy (r(2)  = 0.63). FD also improved the resistance of multifunctionality along the aridity gradient studied. Maintaining and enhancing FD in plant communities may help to buffer negative effects of ongoing global environmental change on dryland multifunctionality.

A global analysis of bidirectional interactions in alpine plant communities shows facilitators experiencing strong reciprocal fitness costs.

Facilitative interactions are defined as positive effects of one species on another, but bidirectional feedbacks may be positive, neutral, or negative. Understanding the bidirectional nature of these interactions is a fundamental prerequisite for the assessment of the potential evolutionary consequences of facilitation. In a global study combining observational and experimental approaches, we quantified the impact of the cover and richness of species associated with alpine cushion plants on reproductive traits of the benefactor cushions. We found a decline in cushion seed production with increasing cover of cushion-associated species, indicating that being a benefactor came at an overall cost. The effect of cushion-associated species was negative for flower density and seed set of cushions, but not for fruit set and seed quality. Richness of cushion-associated species had positive effects on seed density and modulated the effects of their abundance on flower density and fruit set, indicating that the costs and benefits of harboring associated species depend on the composition of the plant assemblage. Our study demonstrates 'parasitic' interactions among plants over a wide range of species and environments in alpine systems, and we consider their implications for the possible selective effects of interactions between benefactor and beneficiary species.

Phenotypic differentiation within a foundation grass species correlates with species richness in a subalpine community.

Few studies have examined consequences of ecotypic differentiation within alpine foundation species for community diversity and their feedbacks for the foundation species' fitness. Additionally, no study has quantified ecotypic differences in competitive effects in the field and in controlled conditions to disentangle genetic from plasticity effects in foundation/subordinate species interactions. We focused on a subalpine community of the French Pyrenees including two phenotypes of a cushion-forming species, Festuca gautieri: tight cushions in dry convex outcrops, and loose cushions (exhibiting high subordinate species richness) in wet concave slopes. We assessed, with field and shadehouse experiments, the genetic vs. plasticity basis of differences in: (1) cushion traits and (2) competitive effects on subordinates, and (3) quantified community feedbacks on foundation species' fitness. We found that trait differences across habitats had both genetic and plasticity bases, with stronger contribution of the latter. Field results showed higher competition within loose than tight phenotypes. In contrast, shadehouse results showed higher competitive ability for tight phenotypes. However, as changes in interactions across habitats were due to environmental effects without changes in cushion effects, we argue that heritable and plastic changes in competitive effects maintain high subordinate species diversity through decreasing competition. We showed high reproduction cost for loose cushions when hosting subordinates highlighting the occurrence of community feedbacks. These results suggest that phenotypic differentiation within foundation species may cascade on subordinate species diversity through heritable and plastic changes in the foundation species' competitive effects, and that community feedbacks may affect foundation species' fitness.

Hotspots of biogeochemical activity linked to aridity and plant traits across global drylands.

Perennial plants create productive and biodiverse hotspots, known as fertile islands, beneath their canopies. These hotspots largely determine the structure and functioning of drylands worldwide. Despite their ubiquity, the factors controlling fertile islands under conditions of contrasting grazing by livestock, the most prevalent land use in drylands, remain virtually unknown. Here we evaluated the relative importance of grazing pressure and herbivore type, climate and plant functional traits on 24 soil physical and chemical attributes that represent proxies of key ecosystem services related to decomposition, soil fertility, and soil and water conservation. To do this, we conducted a standardized global survey of 288 plots at 88 sites in 25 countries worldwide. We show that aridity and plant traits are the major factors associated with the magnitude of plant effects on fertile islands in grazed drylands worldwide. Grazing pressure had little influence on the capacity of plants to support fertile islands. Taller and wider shrubs and grasses supported stronger island effects. Stable and functional soils tended to be linked to species-rich sites with taller plants. Together, our findings dispel the notion that grazing pressure or herbivore type are linked to the formation or intensification of fertile islands in drylands. Rather, our study suggests that changes in aridity, and processes that alter island identity and therefore plant traits, will have marked effects on how perennial plants support and maintain the functioning of drylands in a more arid and grazed world.

Plant biodiversity offsets negative effects of metals and metalloids soil multi-contamination on ecosystem multifunctionality.

Despite increasing metals and metalloids (MM) human-driven soil contamination, how it simultaneously alters biodiversity and ecosystem functioning remains unknown. We used a wide gradient of a 170-year-old MM soil multi-contamination in Mediterranean scrublands to assess the effects of soil multi-contamination on multiple plant biodiversity facets, microbial communities and ecosystem multifunctionality (EMF). We found an overall positive effect of plant biodiversity on EMF mediated by microbial communities, and allowing offsetting the negative impacts of MM soil multi-contamination, especially on soil water holding capacity and nitrogen content. The diversity of distant plant lineages was the key facet promoting EMF by enhancing microbial communities, whereas the subordinate species richness altered EMF. By developing a holistic approach of these complex relationships between soil multi-contamination, plant biodiversity, microbial communities and ecosystem functioning, our results reveal the potential of plant biodiversity, and especially the diversity of evolutionary distant species, to offset the alteration of ecosystem functioning by MM soil multi-contamination. In this worldwide decade of ecosystems restoration, our study helps to identify relevant facets of plant biodiversity promoting contaminated ecosystem functioning, which is crucial to guide and optimize management efforts aiming to restore ecosystems and preserve human health.

Direct and indirect effects of dominant plants on ecosystem multifunctionality.

Biodiversity is essential for the provision of multiple ecosystem functions simultaneously (ecosystem multifunctionality EMF). Yet, it remains unclear whether and how dominant plant species impact EMF. Here, we aimed at disentangling the direct from indirect above- and belowground pathways by which dominant plant species influence EMF. We evaluated the effects of two dominant plant species (Dasiphora fruticosa, and the toxic perennial plant Ligularia virgaurea) with expected positive and negative impacts on the abiotic environment (soil water content and pH), surrounding biological communities (plant and nematode richness, biomass, and abundance in the vicinity), and on the EMF of alpine meadows, respectively. We found that the two dominant plants enhanced EMF, with a positive effect of L. virgaurea on EMF greater than that of D. fruticosa. We also observed that dominant plants impacted on EMF through changes in soil water content and pH (indirect abiotic effects), but not through changes in biodiversity of surrounding plants and nematodes (indirect biotic pathway). Our study suggests that dominant plants may play an important role in promoting EMF, thus expanding the pervasive mass-ratio hypothesis originally framed for individual functions, and could mitigate the negative impacts of vegetation changes on EMF in the alpine meadows.

The interplay of stress and mowing disturbance for the intensity and importance of plant interactions in dry calcareous grasslands.

BACKGROUND AND AIMS: There is still debate regarding the direction and strength of plant interactions under intermediate to high levels of stress. Furthermore, little is known on how disturbance may interact with physical stress in unproductive environments, although recent theory and models have shown that this interplay may induce a collapse of plant interactions and diversity. The few studies assessing such questions have considered the intensity of biotic interactions but not their importance, although this latter concept has been shown to be very useful for understanding the role of interactions in plant communities. The objective of this study was to assess the interplay between stress and disturbance for plant interactions in dry calcareous grasslands. METHODS: A field experiment was set up in the Dordogne, southern France, where the importance and intensity of biotic interactions undergone by four species were measured along a water stress gradient, and with and without mowing disturbance. KEY RESULTS: The importance and intensity of interactions varied in a very similar way along treatments. Under undisturbed conditions, plant interactions switched from competition to neutral with increasing water stress for three of the four species, whereas the fourth species was not subject to any significant biotic interaction along the gradient. Responses to disturbance were more species-specific; for two species, competition disappeared with mowing in the wettest conditions, whereas for the two other species, competition switched to facilitation with mowing. Finally, there were no significant interactions for any species in the disturbed and driest conditions. CONCLUSIONS: At very high levels of stress, plant performances become too weak to allow either competition or facilitation and disturbance may accelerate the collapse of interactions in dry conditions. The results suggest that the importance and direction of interactions are more likely to be positively related in stressful environments.

Grazing and ecosystem service delivery in global drylands.

Grazing represents the most extensive use of land worldwide. Yet its impacts on ecosystem services remain uncertain because pervasive interactions between grazing pressure, climate, soil properties, and biodiversity may occur but have never been addressed simultaneously. Using a standardized survey at 98 sites across six continents, we show that interactions between grazing pressure, climate, soil, and biodiversity are critical to explain the delivery of fundamental ecosystem services across drylands worldwide. Increasing grazing pressure reduced ecosystem service delivery in warmer and species-poor drylands, whereas positive effects of grazing were observed in colder and species-rich areas. Considering interactions between grazing and local abiotic and biotic factors is key for understanding the fate of dryland ecosystems under climate change and increasing human pressure.

Intransitivity increases plant functional diversity by limiting dominance in drylands worldwide.

1. Biotic interactions are key determinants of plant community structure. Indirect interactions such as intransitivity (i.e. in the absence of competitive hierarchies among species) have been hypothesized to benefit diversity within plant communities. However, their effect on functional diversity remains scarcely explored in real communities. Here we develop a novel approach to infer intransitivity from plant spatial patterns and functional traits (height and specific leaf area), and quantify its effect on different components of plant diversity along environmental gradients in 100 drylands from all continents except Antarctica. 2. We first calculated the spatial association pattern for all perennials to infer competition between species. Trait values were used as a proxy of competitive hierarchies to infer the direction of these interactions. We used multiple regression models to evaluate how intransitivity responds to environmental variables (mean annual temperature and precipitation, precipitation seasonality, soil pH, sand content and woody cover). We also used confirmatory path analysis to evaluate the effects of intransitivity on species richness and evenness, trait dispersion and functional diversity. 3. Intransitivity mostly responded to climatic variables, and significantly increased with precipitation scarcity and seasonality. We found that intransitivity had significant effects on functional diversity, mostly by promoting plant community evenness. However, the dominance of woody vegetation (steppes vs. shrublands) modulated this effect. SYNTHESIS: Intransitivity increased the functional diversity of drylands, particularly under high rainfall seasonality, by limiting functionally dominant species. Our findings specify how intransitivity structures the functional diversity of dryland vegetation worldwide. Intransitivity may be particularly important in ecosystems where the availability of abiotic resources changes over time, thereby breaking down inherent competitive hierarchies between plant species. Neglecting intransitivity will bias our estimation of the impacts of biotic interactions on plant communities, a fundamental issue to fully understand how plant communities will respond to ongoing environmental changes.