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Grasslands are integral to maintaining biodiversity and key ecosystem services and are under threat from climate change. Plant and soil microbial diversity, and their interactions, support the provision of multiple ecosystem functions (multifunctionality). However, it remains virtually unknown whether plant and soil microbial diversity explain a unique portion of total variation or shared contributions to supporting multifunctionality across global grasslands. Here, we combine results from a global survey of 101 grasslands with a novel microcosm study, controlling for both plant and soil microbial diversity to identify their individual and interactive contribution to support multifunctionality under aridity and experimental drought. We found that plant and soil microbial diversity independently predict a unique portion of total variation in above- and belowground functioning, suggesting that both types of biodiversity complement each other. Interactions between plant and soil microbial diversity positively impacted multifunctionality including primary production and nutrient storage. Our findings were also climate context dependent, since soil fungal diversity was positively associated with multifunctionality in less arid regions, while plant diversity was strongly and positively linked to multifunctionality in more arid regions. Our results highlight the need to conserve both above- and belowground diversity to sustain grassland multifunctionality in a drier world and indicate climate change may shift the relative contribution of plant and soil biodiversity to multifunctionality across global grasslands., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Martins et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Earth harbours an extraordinary plant phenotypic diversity 1 that is at risk from ongoing global changes 2,3 . However, it remains unknown how increasing aridity and livestock grazing pressure-two major drivers of global change 4-6 -shape the trait covariation that underlies plant phenotypic diversity 1,7 . Here we assessed how covariation among 20 chemical and morphological traits responds to aridity and grazing pressure within global drylands. Our analysis involved 133,769 trait measurements spanning 1,347 observations of 301 perennial plant species surveyed across 326 plots from 6 continents. Crossing an aridity threshold of approximately 0.7 (close to the transition between semi-arid and arid zones) led to an unexpected 88% increase in trait diversity. This threshold appeared in the presence of grazers, and moved toward lower aridity levels with increasing grazing pressure. Moreover, 57% of observed trait diversity occurred only in the most arid and grazed drylands, highlighting the phenotypic uniqueness of these extreme environments. Our work indicates that drylands act as a global reservoir of plant phenotypic diversity and challenge the pervasive view that harsh environmental conditions reduce plant trait diversity 8-10 . They also highlight that many alternative strategies may enable plants to cope with increases in environmental stress induced by climate change and land-use intensification., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Species-specific differences in nutrient acquisition strategies allow for complementary use of resources among plants in mixtures, which may be further shaped by mycorrhizal associations. However, empirical evidence of this potential role of mycorrhizae is scarce, particularly for tree communities. We investigated the impact of tree species richness and mycorrhizal types, arbuscular mycorrhizal fungi (AM) and ectomycorrhizal fungi (EM), on above- and belowground carbon (C), nitrogen (N), and phosphorus (P) dynamics. Soil and soil microbial biomass elemental dynamics showed weak responses to tree species richness and none to mycorrhizal type. However, foliar elemental concentrations, stoichiometry, and pools were significantly affected by both treatments. Tree species richness increased foliar C and P pools but not N pools. Additive partitioning analyses showed that net biodiversity effects of foliar C, N, P pools in EM tree communities were driven by selection effects, but in mixtures of both mycorrhizal types by complementarity effects. Furthermore, increased tree species richness reduced soil nitrate availability, over 2 yr. Our results indicate that positive effects of tree diversity on aboveground nutrient storage are mediated by complementary mycorrhizal strategies and highlight the importance of using mixtures composed of tree species with different types of mycorrhizae to achieve more multifunctional afforestation., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

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.

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 CO 2 modulates future aridity conditions and plant productivity., (© 2021 The Authors New Phytologist © 2021 New Phytologist Foundation.)

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., Competing Interests: The authors declare no competing interest.

Soils harbor a substantial fraction of the world's biodiversity, contributing to many crucial ecosystem functions. It is thus essential to identify general macroecological patterns related to the distribution and functioning of soil organisms to support their conservation and consideration by governance. These macroecological analyses need to represent the diversity of environmental conditions that can be found worldwide. Here we identify and characterize existing environmental gaps in soil taxa and ecosystem functioning data across soil macroecological studies and 17,186 sampling sites across the globe. These data gaps include important spatial, environmental, taxonomic, and functional gaps, and an almost complete absence of temporally explicit data. We also identify the limitations of soil macroecological studies to explore general patterns in soil biodiversity-ecosystem functioning relationships, with only 0.3% of all sampling sites having both information about biodiversity and function, although with different taxonomic groups and functions at each site. Based on this information, we provide clear priorities to support and expand soil macroecological research.

Protists are ubiquitous in soil, where they are key contributors to nutrient cycling and energy transfer. However, protists have received far less attention than other components of the soil microbiome. We used amplicon sequencing of soils from 180 locations across six continents to investigate the ecological preferences of protists and their functional contributions to belowground systems. We complemented these analyses with shotgun metagenomic sequencing of 46 soils to validate the identities of the more abundant protist lineages. We found that most soils are dominated by consumers, although parasites and phototrophs are particularly abundant in tropical and arid ecosystems, respectively. The best predictors of protist composition (primarily annual precipitation) are fundamentally distinct from those shaping bacterial and archaeal communities (namely, soil pH). Some protists and bacteria co-occur globally, highlighting the potential importance of these largely undescribed belowground interactions. Together, this study allowed us to identify the most abundant and ubiquitous protists living in soil, with our work providing a cross-ecosystem perspective on the factors structuring soil protist communities and their likely contributions to soil functioning., (Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. Distributed under a Creative Commons Attribution NonCommercial License 4.0 (CC BY-NC).)

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., Competing Interests: The authors declare no conflict of interest.

Belowground organisms play critical roles in maintaining multiple ecosystem processes, including plant productivity, decomposition, and nutrient cycling. Despite their importance, however, we have a limited understanding of how and why belowground biodiversity (bacteria, fungi, protists, and invertebrates) may change as soils develop over centuries to millennia (pedogenesis). Moreover, it is unclear whether belowground biodiversity changes during pedogenesis are similar to the patterns observed for aboveground plant diversity. Here we evaluated the roles of resource availability, nutrient stoichiometry, and soil abiotic factors in driving belowground biodiversity across 16 soil chronosequences (from centuries to millennia) spanning a wide range of globally distributed ecosystem types. Changes in belowground biodiversity during pedogenesis followed two main patterns. In lower-productivity ecosystems (i.e., drier and colder), increases in belowground biodiversity tracked increases in plant cover. In more productive ecosystems (i.e., wetter and warmer), increased acidification during pedogenesis was associated with declines in belowground biodiversity. Changes in the diversity of bacteria, fungi, protists, and invertebrates with pedogenesis were strongly and positively correlated worldwide, highlighting that belowground biodiversity shares similar ecological drivers as soils and ecosystems develop. In general, temporal changes in aboveground plant diversity and belowground biodiversity were not correlated, challenging the common perception that belowground biodiversity should follow similar patterns to those of plant diversity during ecosystem development. Taken together, our findings provide evidence that ecological patterns in belowground biodiversity are predictable across major globally distributed ecosystem types and suggest that shifts in plant cover and soil acidification during ecosystem development are associated with changes in belowground biodiversity over centuries to millennia., Competing Interests: The authors declare no conflict of interest.

The insurance hypothesis, stating that biodiversity can increase ecosystem stability, has received wide research and political attention. Recent experiments suggest that climate change can impact how plant diversity influences ecosystem stability, but most evidence of the biodiversity-stability relationship obtained to date comes from local studies performed under a limited set of climatic conditions. Here, we investigate how climate mediates the relationships between plant (taxonomical and functional) diversity and ecosystem stability across the globe. To do so, we coupled 14 years of temporal remote sensing measurements of plant biomass with field surveys of diversity in 123 dryland ecosystems from all continents except Antarctica. Across a wide range of climatic and soil conditions, plant species pools, and locations, we were able to explain 73% of variation in ecosystem stability, measured as the ratio of the temporal mean biomass to the SD. The positive role of plant diversity on ecosystem stability was as important as that of climatic and soil factors. However, we also found a strong climate dependency of the biodiversity-ecosystem stability relationship across our global aridity gradient. Our findings suggest that the diversity of leaf traits may drive ecosystem stability at low aridity levels, whereas species richness may have a greater stabilizing role under the most arid conditions evaluated. Our study highlights that to minimize variations in the temporal delivery of ecosystem services related to plant biomass, functional and taxonomic plant diversity should be particularly promoted under low and high aridity conditions, respectively., Competing Interests: The authors declare no conflict of interest.

Recent years have seen a surge of interest in ecosystem multifunctionality, a concept that has developed in the largely separate fields of biodiversity-ecosystem function and land management research. Here we discuss the merit of the multifunctionality concept, the advances it has delivered, the challenges it faces and solutions to these challenges. This involves the redefinition of multifunctionality as a property that exists at two levels: ecosystem function multifunctionality and ecosystem service multifunctionality. The framework presented provides a road map for the development of multifunctionality measures that are robust, quantifiable and relevant to both fundamental ecological science and ecosystem management.

Despite the importance of microbial communities for ecosystem services and human welfare, the relationship between microbial diversity and multiple ecosystem functions and services (that is, multifunctionality) at the global scale has yet to be evaluated. Here we use two independent, large-scale databases with contrasting geographic coverage (from 78 global drylands and from 179 locations across Scotland, respectively), and report that soil microbial diversity positively relates to multifunctionality in terrestrial ecosystems. The direct positive effects of microbial diversity were maintained even when accounting simultaneously for multiple multifunctionality drivers (climate, soil abiotic factors and spatial predictors). Our findings provide empirical evidence that any loss in microbial diversity will likely reduce multifunctionality, negatively impacting the provision of services such as climate regulation, soil fertility and food and fibre production by terrestrial ecosystems.

Intransitive competition networks, those in which there is no single best competitor, may ensure species coexistence. However, their frequency and importance in maintaining diversity in real-world ecosystems remain unclear. We used two large data sets from drylands and agricultural grasslands to assess: (1) the generality of intransitive competition, (2) intransitivity-richness relationships and (3) effects of two major drivers of biodiversity loss (aridity and land-use intensification) on intransitivity and species richness. Intransitive competition occurred in > 65% of sites and was associated with higher species richness. Intransitivity increased with aridity, partly buffering its negative effects on diversity, but was decreased by intensive land use, enhancing its negative effects on diversity. These contrasting responses likely arise because intransitivity is promoted by temporal heterogeneity, which is enhanced by aridity but may decline with land-use intensity. We show that intransitivity is widespread in nature and increases diversity, but it can be lost with environmental homogenisation., (© 2015 John Wiley & Sons Ltd/CNRS.)

Once seen as anomalous, facilitative interactions among plants and their importance for community structure and functioning are now widely recognized. The growing body of modelling, descriptive and experimental studies on facilitation covers a wide variety of terrestrial and aquatic systems throughout the globe. However, the lack of a general body of theory linking facilitation among different types of organisms and biomes and their responses to environmental changes prevents further advances in our knowledge regarding the evolutionary and ecological implications of facilitation in plant communities. Moreover, insights gathered from alternative lines of inquiry may substantially improve our understanding of facilitation, but these have been largely neglected thus far. Despite over 15 years of research and debate on this topic, there is no consensus on the degree to which plant-plant interactions change predictably along environmental gradients (i.e. the stress-gradient hypothesis), and this hinders our ability to predict how plant-plant interactions may affect the response of plant communities to ongoing global environmental change. The existing controversies regarding the response of plant-plant interactions across environmental gradients can be reconciled when clearly considering and determining the species-specificity of the response, the functional or individual stress type, and the scale of interest (pairwise interactions or community-level response). Here, we introduce a theoretical framework to do this, supported by multiple lines of empirical evidence. We also discuss current gaps in our knowledge regarding how plant-plant interactions change along environmental gradients. These include the existence of thresholds in the amount of species-specific stress that a benefactor can alleviate, the linearity or non-linearity of the response of pairwise interactions across distance from the ecological optimum of the beneficiary, and the need to explore further how frequent interactions among multiple species are and how they change across different environments. We review the latest advances in these topics and provide new approaches to fill current gaps in our knowledge. We also apply our theoretical framework to advance our knowledge on the evolutionary aspects of plant facilitation, and the relative importance of facilitation, in comparison with other ecological processes, for maintaining ecosystem structure, functioning and dynamics. We build links between these topics and related fields, such as ecological restoration, woody encroachment, invasion ecology, ecological modelling and biodiversity-ecosystem-functioning relationships. By identifying commonalities and insights from alternative lines of research, we further advance our understanding of facilitation and provide testable hypotheses regarding the role of (positive) biotic interactions in the maintenance of biodiversity and the response of ecological communities to ongoing environmental changes., (© 2014 The Authors. Biological Reviews © 2014 Cambridge Philosophical Society.)

Drought is an increasingly common phenomenon in drylands as a consequence of climate change. We used 311 sites across a broad range of environmental conditions in Patagonian rangelands to evaluate how drought severity and temperature (abiotic factors) and vegetation structure (biotic factors) modulate the impact of a drought event on the annual integral of normalized difference vegetation index (NDVI-I), our surrogate of ecosystem functioning. We found that NDVI-I decreases were larger with both increasing drought severity and temperature. Plant species richness (SR) and shrub cover (SC) attenuated the effects of drought on NDVI-I. Grass cover did not affect the impacts of drought on NDVI-I. Our results suggest that warming and species loss, two important imprints of global environmental change, could increase the vulnerability of Patagonian ecosystems to drought. Therefore, maintaining SR through appropriate grazing management can attenuate the adverse effects of climate change on ecosystem functioning., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

Dryland ecosystems account for ca. 27% of global soil organic carbon (C) reserves, yet it is largely unknown how climate change will impact C cycling and storage in these areas. In drylands, soil C concentrates at the surface, making it particularly sensitive to the activity of organisms inhabiting the soil uppermost levels, such as communities dominated by lichens, mosses, bacteria and fungi (biocrusts). We conducted a full factorial warming and rainfall exclusion experiment at two semiarid sites in Spain to show how an average increase of air temperature of 2-3 °C promoted a drastic reduction in biocrust cover (ca. 44% in 4 years). Warming significantly increased soil CO2 efflux, and reduced soil net CO2 uptake, in biocrust-dominated microsites. Losses of biocrust cover with warming through time were paralleled by increases in recalcitrant C sources, such as aromatic compounds, and in the abundance of fungi relative to bacteria. The dramatic reduction in biocrust cover with warming will lessen the capacity of drylands to sequester atmospheric CO2 . This decrease may act synergistically with other warming-induced effects, such as the increase in soil CO2 efflux and the changes in microbial communities to alter C cycling in drylands, and to reduce soil C stocks in the mid to long term., (© 2013 John Wiley & Sons Ltd.)

Experiments suggest that biodiversity enhances the ability of ecosystems to maintain multiple functions, such as carbon storage, productivity, and the buildup of nutrient pools (multifunctionality). However, the relationship between biodiversity and multifunctionality has never been assessed globally in natural ecosystems. We report here on a global empirical study relating plant species richness and abiotic factors to multifunctionality in drylands, which collectively cover 41% of Earth's land surface and support over 38% of the human population. Multifunctionality was positively and significantly related to species richness. The best-fitting models accounted for over 55% of the variation in multifunctionality and always included species richness as a predictor variable. Our results suggest that the preservation of plant biodiversity is crucial to buffer negative effects of climate change and desertification in drylands.

The study of species coexistence and community assembly has been a hot topic in ecology for decades. Disentangling the hierarchical role of abiotic and biotic filters is crucial to understand community assembly processes. The most critical environmental factor in semi-arid environments is known to be water availability, and perennials are usually described as nurses that create milder local conditions and expand the niche range of several species. We aimed to broaden this view by jointly evaluating how biological soil crusts (BSCs), water availability, perennial species (presence/absence of Stipa tenacissima) and plant-plant interactions shape a semi-arid annual plant community. The presence and cover of annual species was monitored during three years of contrasting climate. Water stress acted as the primary filter determining the species pool available for plant community assembly. Stipa and BSCs acted as secondary filters by modulating the effects of water availability. At extremely harsh environmental conditions, Stipa exerted a negative effect on the annual plant community, while at more benign conditions it increased annual community richness. Biological soil crusts exerted a contradictory effect depending on climate and on the presence of Stipa, favoring annuals in the most adverse conditions but showing repulsion at higher water availability conditions. Finally, interactions among co-occurring annuals shaped species richness and diversity of the final annual plant assembly. This study sheds light on the processes determining the assembly of annual communities and highlights the importance of Biological Soil Crusts and of interactions among annual plants on the final outcome of the species assembly.

Climate change will exacerbate the degree of abiotic stress experienced by semi-arid ecosystems. While abiotic stress profoundly affects biotic interactions, their potential role as modulators of ecosystem responses to climate change is largely unknown. Using plants and biological soil crusts, we tested the relative importance of facilitative-competitive interactions and other community attributes (cover, species richness and species evenness) as drivers of ecosystem functioning along stress gradients in semi-arid Mediterranean ecosystems. Biotic interactions shifted from facilitation to competition along stress gradients driven by water availability and temperature. These changes were, however, dependent on the spatial scale and the community considered. We found little evidence to suggest that biotic interactions are a major direct influence upon indicators of ecosystem functioning (soil respiration, organic carbon, water-holding capacity, compaction and the activity of enzymes related to the carbon, nitrogen and phosphorus cycles) along stress gradients. However, attributes such as cover and species richness showed a direct effect on ecosystem functioning. Our results do not agree with predictions emphasizing that the importance of plant-plant interactions will be increased under climate change in dry environments, and indicate that reductions in the cover of plant and biological soil crust communities will negatively impact ecosystems under future climatic conditions.

Belowground organisms play critical roles in maintaining multiple ecosystem processes, including plant productivity, decomposition, and nutrient cycling. Despite their importance, however, we have a limited understanding of how and why belowground biodiversity (bacteria, fungi, protists, and invertebrates) may change as soils develop over centuries to millennia (pedogenesis). Moreover, it is unclear whether belowground biodiversity changes during pedogenesis are similar to the patterns observed for aboveground plant diversity. Here we evaluated the roles of resource availability, nutrient stoichiometry, and soil abiotic factors in driving belowground biodiversity across 16 soil chronosequences (from centuries to millennia) spanning a wide range of globally distributed ecosystem types. Changes in belowground biodiversity during pedogenesis followed two main patterns. In lower-productivity ecosystems (i.e., drier and colder), increases in belowground biodiversity tracked increases in plant cover. In more productive ecosystems (i.e., wetter and warmer), increased acidification during pedogenesis was associated with declines in belowground biodiversity. Changes in the diversity of bacteria, fungi, protists, and invertebrates with pedogenesis were strongly and positively correlated worldwide, highlighting that belowground biodiversity shares similar ecological drivers as soils and ecosystems develop. In general, temporal changes in aboveground plant diversity and belowground biodiversity were not correlated, challenging the common perception that belowground biodiversity should follow similar patterns to those of plant diversity during ecosystem development. Taken together, our findings provide evidence that ecological patterns in belowground biodiversity are predictable across major globally distributed ecosystem types and suggest that shifts in plant cover and soil acidification during ecosystem development are associated with changes in belowground biodiversity over centuries to millennia.

The emergence of high-throughput DNA sequencing methods provides unprecedented opportunities to further unravel bacterial biodiversity and its worldwide role from human health to ecosystem functioning. However, despite the abundance of sequencing studies, combining data from multiple individual studies to address macroecological questions of bacterial diversity remains methodically challenging and plagued with biases. Here, using a machine-learning approach that accounts for differences among studies and complex interactions among taxa, we merge 30 independent bacterial data sets comprising 1,998 soil samples from 21 countries. Whereas previous meta-analysis efforts have focused on bacterial diversity measures or abundances of major taxa, we show that disparate amplicon sequence data can be combined at the taxonomy-based level to assess bacterial community structure. We find that rarer taxa are more important for structuring soil communities than abundant taxa, and that these rarer taxa are better predictors of community structure than environmental factors, which are often confounded across studies. We conclude that combining data from independent studies can be used to explore bacterial community dynamics, identify potential 'indicator' taxa with an important role in structuring communities, and propose hypotheses on the factors that shape bacterial biogeography that have been overlooked in the past.

Fungi are highly diverse organisms, which provide multiple ecosystem services. However, compared with charismatic animals and plants, the distribution patterns and conservation needs of fungi have been little explored. Here, we examined endemicity patterns, global change vulnerability and conservation priority areas for functional groups of soil fungi based on six global surveys using a high‐resolution, long‐read metabarcoding approach. We found that the endemicity of all fungi and most functional groups peaks in tropical habitats, including Amazonia, Yucatan, West‐Central Africa, Sri Lanka, and New Caledonia, with a negligible island effect compared with plants and animals. We also found that fungi are predominantly vulnerable to drought, heat and land‐cover change, particularly in dry tropical regions with high human population density. Fungal conservation areas of highest priority include herbaceous wetlands, tropical forests, and woodlands. We stress that more attention should be focused on the conservation of fungi, especially root symbiotic arbuscular mycorrhizal and ectomycorrhizal fungi in tropical regions as well as unicellular early‐diverging groups and macrofungi in general. Given the low overlap between the endemicity of fungi and macroorganisms, but high conservation needs in both groups, detailed analyses on distribution and conservation requirements are warranted for other microorganisms and soil organisms. [ABSTRACT FROM AUTHOR]

In recent years there has been an upsurge of studies on ecosystem multifunctionality (EMF), or the ability of ecosystems to simultaneously provide multiple functions and/or services. The concept of EMF itself, the analytical approaches used to calculate it, and its implications depending on the spatial scale and field of study have been discussed in detail. However, to date there has been little dialogue concerning the basis of EMF studies: what should or should not be considered appropriate measures for ecosystem functions.To begin this discussion, we performed an in‐depth review of EMF studies across four major terrestrial ecosystems (agroecosystems, drylands, forests and grasslands) by analysing 82 studies, which together have assessed 775 ecosystem functions from a variety of field and greenhouse experiments across the globe.The number of ecosystem functions analysed varied from two to 82 per study and we found large differences in the distribution of functions across ecosystem types and ecosystem service categories. Furthermore, there was little explanation of why certain variables were included in the EMF calculation or how they relate to ecosystem functioning.Synthesis. Based on the literature analysis, it is clear that there is no general agreement regarding which measurements should or should not be considered functions in the field of ecology. To address this issue, we propose a general guideline for determining and measuring appropriate functions. [ABSTRACT FROM AUTHOR]

Aims Biodiversity is often positively related to the capacity of an ecosystem to provide multiple functions simultaneously (i.e. multifunctionality). However, there is some controversy over whether biodiversity–multifunctionality relationships depend on the number of functions considered. Particularly, investigators have documented contrasting findings that the effects of biodiversity on ecosystem multifunctionality do not change or increase with the number of ecosystem functions. Here, we provide some clarity on this issue by examining the statistical underpinnings of different multifunctionality metrics. Methods We used simulations and data from a variety of empirical studies conducted across spatial scales (from local to global) and biomes (temperate and alpine grasslands, forests and drylands). We revisited three methods to quantify multifunctionality including the averaging approach, summing approach and threshold-based approach. Important Findings Biodiversity–multifunctionality relationships either did not change or increased as more functions were considered. These results were best explained by the statistical underpinnings of the averaging and summing multifunctionality metrics. Specifically, by averaging the individual ecosystem functions, the biodiversity–multifunctionality relationships equal the population mean of biodiversity-single function relationships, and thus will not change with the number of functions. Likewise, by summing the individual ecosystem functions, the strength of biodiversity–multifunctionality relationships increases as the number of functions increased. We proposed a scaling standardization method by converting the averaging or summing metrics into a scaling metric, which would make comparisons among different biodiversity studies. In addition, we showed that the range-relevant standardization can be applied to the threshold-based approach by solving for the mathematical artefact of the approach (i.e. the effects of biodiversity may artificially increase with the number of functions considered). Our study highlights different approaches yield different results and that it is essential to develop an understanding of the statistical underpinnings of different approaches. The standardization methods provide a prospective way of comparing biodiversity–multifunctionality relationships across studies. [ABSTRACT FROM AUTHOR]

Past vegetation and climatic conditions are known to influence current biodiversity patterns. However, whether their legacy effects affect the provision of multiple ecosystem functions, that is, multifunctionality, remains largely unknown. Here we analyzed soil nutrient stocks and their transformation rates in 236 drylands from six continents to evaluate the associations between current levels of multifunctionality and legacy effects of the Last Glacial Maximum (LGM) desert biome distribution and climate. We found that past desert distribution and temperature legacy, defined as increasing temperature from LGM, were negatively correlated with contemporary multifunctionality even after accounting for predictors such as current climate, soil texture, plant species richness, and site topography. Ecosystems that have been deserts since the LGM had up to 30% lower contemporary multifunctionality compared with those that were nondeserts during the LGM. In addition, ecosystems that experienced higher warming rates since the LGM had lower contemporary multifunctionality than those suffering lower warming rates, with a ~9% reduction per extra degree Celsius. Past desert distribution and temperature legacies had direct negative effects, while temperature legacy also had indirect (via soil sand content) negative effects on multifunctionality. Our results indicate that past biome and climatic conditions have left a strong "functionality debt" in global drylands. They also suggest that ongoing warming and expansion of desert areas may leave a strong fingerprint in the future functioning of dryland ecosystems worldwide that needs to be considered when establishing management actions aiming to combat land degradation and desertification. [ABSTRACT FROM AUTHOR]

Aim: Species–area relationships (also known as "species–area curves" and "species accumulation curves") represent the relationship between species richness and the area sampled in a given community. These relationships can be used to describe diversity patterns while accounting for the well‐known scale‐dependence of species richness. Despite their value, their functional form and parameters, as well as their determinants, have barely been investigated in drylands. Location: 171 drylands from all continents except Antarctica. Time period: 2006–2013. Major taxa studied: Perennial plants. Methods: We characterized species–area relationships of plant communities by building accumulation curves describing the expected number of species as a function of the number of sampling units, and later compared the fit of three functions (power law, logarithmic and Michaelis–Menten). We tested the prediction that the effects of aridity, soil pH on the species–area relationship (SAR) are mediated by vegetation attributes such as evenness, cover and spatial aggregation. Results: We found that the logarithmic relationship was the most common functional form (c. 50%), followed by Michaelis–Menten (c. 33%) and power law (c. 17%). Functional form was mainly determined by evenness. Power‐law relationships were found mostly under low evenness, logarithmic relationships peaked under intermediate evenness and the Michaelis–Menten function increased in frequency with increasing evenness. The SAR parameters approximated by the logarithmic model ["small‐scale richness" (b0) and "accumulation coefficient" (b1)] were determined by vegetation attributes. Increasing spatial aggregation had a negative effect on the small‐scale richness and a positive effect on the accumulation coefficient, while evenness had an opposite effect. In addition, the accumulation coefficient was positively affected by cover. Interestingly, increasing aridity decreased small‐scale richness but did not affect the accumulation coefficient. Main conclusions: Our findings highlight the role of evenness, spatial aggregation and cover as the main drivers of SARs in drylands, the Earth's largest biome. [ABSTRACT FROM AUTHOR]

Despite their importance, how plant communities and soil microorganisms interact to determine the capacity of ecosystems to provide multiple functions simultaneously (multifunctionality) under climate change is poorly known. We conducted a common garden experiment using grassland species to evaluate how plant functional structure and soil microbial (bacteria and protists) diversity and abundance regulate soil multifunctionality responses to joint changes in plant species richness (one, three and six species) and simulated climate change (3°C warming and 35% rainfall reduction). The effects of species richness and climate on soil multifunctionality were indirectly driven via changes in plant functional structure and their relationships with the abundance and diversity of soil bacteria and protists. More specifically, warming selected for the larger and most productive plant species, increasing the average size within communities and leading to reductions in functional plant diversity. These changes increased the total abundance of bacteria that, in turn, increased that of protists, ultimately promoting soil multifunctionality. Our work suggests that cascading effects between plant functional traits and the abundance of multitrophic soil organisms largely regulate the response of soil multifunctionality to simulated climate change, and ultimately provides novel experimental insights into the mechanisms underlying the effects of biodiversity and climate change on ecosystem functioning. How plant communities and soil microorganisms interact to determine the multifunctionality under climate change is poorly known. We evaluated, using an experimental approach, how simulated climate change (warming and rainfall reduction) and initial plant species richness affect plant functional structure and soil microorganisms. Also, we assessed how all these factors affect, directly and indirectly, soil multifunctionality. Warming selected for the larger and most productive plant species, increasing the average size in the communities and leading to reductions in functional plant diversity. These changes increased the total abundance of bacteria that, in turn, increased that of protists, ultimately promoting soil multifunctionality. [ABSTRACT FROM AUTHOR]

Summary: Recent research indicates that increased aridity linked to climate change will reduce the diversity of soil microbial communities and shift their community composition in drylands, Earth's largest biome. However, we lack both a theoretical framework and solid empirical evidence of how important biotic components from drylands, such as biocrust‐forming mosses, will regulate the responses of microbial communities to expected increases in aridity with climate change.Here we report results from a cross‐continental (North America, Europe and Australia) survey of 39 locations from arid to humid ecosystems, where we evaluated how biocrust‐forming mosses regulate the relationship between aridity and the community composition and diversity of soil bacteria and fungi in dryland ecosystems.Increasing aridity was negatively related to the richness of fungi, and either positively or negatively related to the relative abundance of selected microbial phyla, when biocrust‐forming mosses were absent. Conversely, we found an overall lack of relationship between aridity and the relative abundance and richness of microbial communities under biocrust‐forming mosses.Our results suggest that biocrust‐forming mosses mitigate the impact of aridity on the community composition of globally distributed microbial taxa, and the diversity of fungi. They emphasize the importance of maintaining biocrusts as a sanctuary for soil microbes in drylands. [ABSTRACT FROM AUTHOR]

Understanding how drylands respond to ongoing environmental change is extremely important for global sustainability. In this review, we discuss how biotic attributes, climate, grazing pressure, land cover change, and nitrogen deposition affect the functioning of drylands at multiple spatial scales. Our synthesis highlights the importance of biotic attributes (e.g., species richness) in maintaining fundamental ecosystem processes such as primary productivity, illustrates how nitrogen deposition and grazing pressure are impacting ecosystem functioning in drylands worldwide, and highlights the importance of the traits of woody species as drivers of their expansion in former grasslands. We also emphasize the role of attributes such as species richness and abundance in controlling the responses of ecosystem functioning to climate change. This knowledge is essential to guide conservation and restoration efforts in drylands, as biotic attributes can be actively managed at the local scale to increase ecosystem resilience to global change. [ABSTRACT FROM AUTHOR]

Despite the vital role of microorganisms for ecosystem functioning and human welfare, our understanding of their global diversity and biogeographical patterns lags significantly behind that of plants and animals. We conducted a meta-analysis including ~600 soil samples from all continents to evaluate the biogeographical patterns and drivers of bacterial diversity in terrestrial ecosystems at the global scale. Similar to what has been found with plants and animals, the diversity of soil bacteria in the Southern Hemisphere decreased from the equator to Antarctica. However, soil bacteria showed similar levels of diversity across the Northern Hemisphere. The composition of bacterial communities followed dissimilar patterns between hemispheres, as the Southern and Northern Hemispheres were dominated by Actinobacteria and Acidobacteria, respectively. However, Proteobacteria was co-dominant in both hemispheres. Moreover, we found a decrease in soil bacterial diversity with altitude. Climatic features (e.g., high diurnal temperature range and low temperature) were correlated with the lower diversity found at high elevations, but geographical gradients in soil total carbon and species turnover were important drivers of the observed latitudinal patterns. We thus found both parallels and differences in the biogeographical patterns of aboveground vs. soil bacterial diversity. Our findings support previous studies that highlighted soil pH, spatial influence, and organic matter as important drivers of bacterial diversity and composition. Furthermore, our results provide a novel integrative view of how climate and soil factors influence soil bacterial diversity at the global scale, which is critical to improve ecosystem and earth system simulation models and for formulating sustainable ecosystem management and conservation policies. [ABSTRACT FROM AUTHOR]

Positive interactions among plants can increase species richness by relaxing environmental filters and providing more heterogeneous environments. However, it is not known if facilitation could affect coexistence through other mechanisms. Most studies on plant coexistence focus on negative frequency-dependent mechanisms (decreasing the abundance of common species); here, we test if facilitation can enhance coexistence by giving species an advantage when rare., To test our hypothesis, we used a global data set from drylands and alpine environments and measured the intensity of facilitation (based on co-occurrences with nurse plants) for 48 species present in at least 4 different sites and with a range of abundances in the field. We compared these results with the degree of facilitation experienced by species which are globally rare or common (according to the IUCN Red List), and with a larger data base including over 1200 co-occurrences of target species with their nurses., Facilitation was stronger for rare species (i.e. those having lower local abundances or considered endangered by the IUCN) than for common species, and strongly decreased with the abundance of the facilitated species. These results hold after accounting for the distance of each species from its ecological optimum (i.e. the degree of functional stress it experiences)., Synthesis. Our results highlight that nurse plants not only increase the number of species able to colonize a given site, but may also promote species coexistence by preventing the local extinction of rare species. Our findings illustrate the role that nurse plants play in conserving endangered species and link the relationship between facilitation and diversity with coexistence theory. As such, they provide further mechanistic understanding on how facilitation maintains plant diversity. [ABSTRACT FROM AUTHOR]

Background and aims: Many previous studies have evaluated aboveground-heterotrophic belowground interactions such as plant-soil feedbacks, plant-mycorrhizal fungi associations or plant-actinorhizal symbioses. However, few studies have used biocrusts, which are specialized soil communities of autotrophic cyanobacteria, mosses, lichens and non-photosynthetic fungi and bacteria that are prevalent in drylands worldwide. These communities largely influence ecosystem functioning, and can be used as a model system for studying above-belowground interactions. In this study, we evaluated how biocrusts affect the functional diversity and biomass of microbial diversities beneath biocrusts. Methods: We performed two microcosm experiments using biocrust-forming lichens where we manipulated their biotic attributes to test independently the effects of species richness (from two to eight species), composition, evenness (maximal and low evenness) and spatial pattern (clumped and random distribution) on the microbial catabolic profile and microbial functional diversity. Results: Microcosms with a random pattern had a higher microbial catabolic profile than those with a clumped pattern. Significant richness × evenness × pattern and richness × evenness interactions were found when analyzing microbial catabolic profile and biomass, respectively. Microcosms with a random pattern, intermediate number of species, and maximal evenness level had higher microbial catabolic profile. At the maximal evenness level, assemblages had higher microbial catabolic profile and microbial biomass when they contained four species. The richness × evenness × pattern interaction was the most informative predictor of variations in microbial catabolic profile. Conclusions: Our results indicate that soil microorganisms are influenced by biocrusts, just as they are influenced by plants, and highlight the importance of higher order interactions among species richness, evenness, and spatial pattern as drivers of microbial communities. The results also emphasize the importance of studying several biotic attributes simultaneously when studying biocrust-soil microorganism interactions, as in nature, community properties do not exert their influence in isolation. [ABSTRACT FROM AUTHOR]

In a network of competing species, a competitive intransitivity occurs when the ranking of competitive abilities does not follow a linear hierarchy (A > B > C but C > A). A variety of mathematical models suggests that intransitive networks can prevent or slow down competitive exclusion and maintain biodiversity by enhancing species coexistence. However, it has been difficult to assess empirically the relative importance of intransitive competition because a large number of pairwise species competition experiments are needed to construct a competition matrix that is used to parameterize existing models. Here we introduce a statistical framework for evaluating the contribution of intransitivity to community structure using species abundance matrices that are commonly generated from replicated sampling of species assemblages. We provide metrics and analytical methods for using abundance matrices to estimate species competition and patch transition matrices by using reverse-engineering and a colonization-competition model. These matrices provide complementary metrics to estimate the degree of intransitivity in the competition network of the sampled communities. Benchmark tests reveal that the proposed methods could successfully detect intransitive competition networks, even in the absence of direct measures of pairwise competitive strength. To illustrate the approach, we analyzed patterns of abundance and biomass of five species of necrophagous Diptera and eight species of their hymenopteran parasitoids that co-occur in beech forests in Germany. We found evidence for a strong competitive hierarchy within communities of flies and parasitoids. However, for parasitoids, there was a tendency towards increasing intransitivity in higher weight classes, which represented larger resource patches. These tests provide novel methods for empirically estimating the degree of intransitivity in competitive networks from observational datasets. They can be applied to experimental measures of pairwise species interactions, as well as to spatio-temporal samples of assemblages in homogenous environments or environmental gradients. [ABSTRACT FROM AUTHOR]

Habitat filtering ( HF, trait convergence) and niche differentiation ( ND, trait divergence) are known to impact upon plant community structure. Both processes integrate individual responses to the abiotic environment and biotic interactions. Thus, it is difficult to clearly identify the underlying abiotic and biotic factors that ultimately impact community structure by looking at community-level patterns of trait divergence or convergence alone., We used a functional trait-based and multiscale approach to assess how biotic interactions and aridity determine the functional structure of semi-arid shrublands sampled along a large aridity gradient in Spain. At the regional scale, we investigated functional differences among species (axes of specialization) to identify important traits for community assembly. At the community scale, we evaluated the relative impact of HF and ND on community structure using a null model approach. Finally, at the plant neighbourhood scale, we evaluated the impact of biotic interactions on community structure by investigating the spatial patterns of trait aggregation., The shrub species surveyed can be separated along four axes of specialization based on their above-ground architecture and leaf morphology. Our community scale analysis suggested that the functional structure of semi-arid communities was clearly non-random, HF and ND acting independently on different traits to determine community structure along the aridity gradient. At the plant neighbourhood scale, the spatial distribution of species was also clearly not random, suggesting that competition and facilitation impacted on the observed changes in the functional diversity of shrubland communities along the aridity gradient., Synthesis: Our results demonstrated that HF and ND acted simultaneously on independent traits to jointly determine community structure. Most importantly, our multiscale approach suggested that competition and facilitation interplayed with aridity to determine this structure. Competition appeared to be constant along the aridity gradient and explained the high functional diversity observed in semi-arid shrublands. Facilitation affected subordinate and rare species and, thus, may act to enhance the biodiversity of these ecosystems. Finally, the framework employed in our study allows moving forward from the examination of patterns to the development of mechanistic trait-based approaches to study plant community assembly. [ABSTRACT FROM AUTHOR]

Recent research has shown that biodiversity may have its greatest impact on ecosystem functioning in heterogeneous environments. However, the role of soil heterogeneity as a modulator of ecosystem responses to changes in biodiversity remains poorly understood, as few biodiversity studies have explicitly considered this important ecosystem feature. We conducted a microcosm experiment over two growing seasons to evaluate the joint effects of changes in plant functional groups (grasses, legumes, non-legume forbs and a combination of them), spatial distribution of soil nutrients (homogeneous and heterogeneous) and nutrient availability (50 and 100 mg of nitrogen (N) added as organic material) on plant productivity and surrogates of carbon, phosphorous and N cycling (β-glucosidase and acid phosphatase enzymes and in situ N availability, respectively). Soil nutrient heterogeneity interacted with nutrient availability and plant functional diversity to determine productivity and nutrient cycling responses. All the functional groups exhibited precise root foraging patterns. Above- and below-ground productivity increased under heterogeneous nutrient supply. Surrogates of nutrient cycling were not directly affected by soil nutrient heterogeneity. Regardless of their above- and below-ground biomass, legumes increased the availability of soil inorganic N and the activity of the acid phosphatase and β-glucosidase enzymes. Our study emphasizes the role of soil nutrient heterogeneity as a modulator of ecosystem responses to changes in functional diversity beyond the species level. Functional group identity, rather than richness, can play a key role in determining the effects of biodiversity on ecosystem functioning. Synthesis. Our results highlight the importance of explicitly considering soil heterogeneity in diversity-ecosystem functioning experiments, where the identity of the plant functional group is of major importance. Such consideration will improve our ability to fully understand the role of plant diversity on ecosystem functioning in ubiquitous heterogeneous environments. [ABSTRACT FROM AUTHOR]

Abstract: Despite that soils may be the greatest repository of biodiversity on Earth, and that most terrestrial ecosystem functions occur in the soil, research on the role of soil biodiversity in ecosystem function has lagged behind corresponding research on aboveground organisms. Soil organisms pose special problems to biodiversity–function research, including the fact that we usually do not know their identity nor what they do in soil ecosystems, cannot easily estimate their biodiversity, and cannot culture the majority of the organisms for use in manipulative experiments. We propose here that biological soil crusts (BSCs) of deserts and many other ecosystems may serve as a useful model system for diversity–function research because the species concept is relatively well-defined within BSC organisms, their functional attributes are relatively well-known, and estimation and manipulation of biodiversity in experiments are feasible, at least within some groups of BSC biota. In spite of these features, there is a pronounced lack of research on biodiversity–function using these organisms. At least two complementary approaches are possible: experiments using artificially-constructed BSCs, and observational studies which statistically control for the effects of other factors which are likely to covary with biodiversity. We applied the latter to four observational datasets collected at multiple spatial scales in Spain and the United States using structural equation models or path analysis using ecosystem function indicators relating to hydrology, trapping and retention of soil resources, and nutrient cycling. We found that, even when total BSC abundance and key environmental gradients are controlled for, direct and approximately linear relationships between species richness and/or evenness and indicators of ecosystem functioning were common. Such relationships appear to vary independently of region or spatial scale, but their strength seems to differ in every dataset. Functional group richness did not seem to adequately capture biodiversity–function relationships, suggesting that bryophyte and lichen components of BSC may exhibit low redundancy. More research employing the multi-trophic, multi-functional, and manipulable BSC system may enable more rapid understanding of the consequences of biodiversity loss in soils, and help enable a biodiversity–function theory that is pertinent to the numerous ecosystem services provided by soil organisms. [Copyright &y& Elsevier]

Changes in the atmospheric concentration of carbon dioxide ([CO2]), nutrient availability and biotic diversity are three major drivers of the ongoing global change impacting terrestrial ecosystems worldwide. While it is well established that soil nutrient heterogeneity exerts a strong influence on the development of plant individuals and communities, it is virtually unknown how nutrient heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. We conducted a microcosm experiment to evaluate the effect of simultaneous changes in [CO2], nutrient heterogeneity (NH), nutrient availability (NA) and species evenness on the biomass and nutrient uptake patterns of assemblages formed by Lolium perenne, Plantago lanceolata and Holcus lanatus. When the nutrients were heterogeneously supplied, assemblages exhibited precise root foraging patterns, and had higher above- and belowground biomass (average increases of 32% and 29% for above- and belowground biomass, respectively). Nutrient heterogeneity also modulated the effects of NA on biomass production, complementarity in nitrogen uptake and below: aboveground ratio, as well as those of [CO2] on the nutrient use efficiency at the assemblage level. Our results show that nutrient heterogeneity has the potential to influence the response of plant assemblages to simultaneous changes in [CO2], nutrient availability and biotic diversity, and suggest that it is an important environmental factor to interpret and assess plant assemblage responses to global change. [ABSTRACT FROM AUTHOR]

Summary: A direct relationship between ecosystem structure and function has been widely accepted by restoration ecologists. According to this paradigm, ecosystem degradation and aggradation represent parallel changes in structure and function, restoration following the same path as spontaneous succession. But the existence of single bidirectional trajectories and endpoints is not supported by empirical evidence. On the contrary, multiple meta-stable states, irreversible changes and hysteresis are common in nature. These situations are better described by state-and-transition models. Merging those models into the structure–function framework may help to develop new hypotheses on ecosystem dynamics, and may provide a suitable framework for planning restoration activities. We use the relationship between ecosystem function and the effort needed to restore a degraded ecosystem (i.e. restorability) as an example. A linear relationship between ecosystem structure and function suggests that ecosystem degradation and restorability are directly related. This may not be true when multiple states, not necessarily connected, are considered. We show two case studies that support this point, and discuss the implications of the incorporation of state-and-transition models into the structure–function framework on relevant topics of restoration ecology and conservation biology, such as the choice of reference ecosystems, the evaluation of restoration actions, and the identification of priority areas for conservation and restoration. [Copyright &y& Elsevier]