Search

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought., Competing Interests: Competing interests statement:The authors declare no competing interest.

Soil fungi are a key constituent of global biodiversity and play a pivotal role in agroecosystems. How arable farming affects soil fungal biogeography and whether it has a disproportional impact on rare taxa is poorly understood. Here, we used the high-resolution PacBio Sequel targeting the entire ITS region to investigate the distribution of soil fungi in 217 sites across a 3000 km gradient in Europe. We found a consistently lower diversity of fungi in arable lands than grasslands, with geographic locations significantly impacting fungal community structures. Prevalent fungal groups became even more abundant, whereas rare groups became fewer or absent in arable lands, suggesting a biotic homogenization due to arable farming. The rare fungal groups were narrowly distributed and more common in grasslands. Our findings suggest that rare soil fungi are disproportionally affected by arable farming, and sustainable farming practices should protect rare taxa and the ecosystem services they support., (© 2024. The Author(s).)

How the multiple facets of soil fungal diversity vary worldwide remains virtually unknown, hindering the management of this essential species-rich group. By sequencing high-resolution DNA markers in over 4000 topsoil samples from natural and human-altered ecosystems across all continents, we illustrate the distributions and drivers of different levels of taxonomic and phylogenetic diversity of fungi and their ecological groups. We show the impact of precipitation and temperature interactions on local fungal species richness (alpha diversity) across different climates. Our findings reveal how temperature drives fungal compositional turnover (beta diversity) and phylogenetic diversity, linking them with regional species richness (gamma diversity). We integrate fungi into the principles of global biodiversity distribution and present detailed maps for biodiversity conservation and modeling of global ecological processes.

Soils support an immense portion of Earth's biodiversity and maintain multiple ecosystem functions which are essential for human well-being. Environmental thresholds are known to govern global vegetation patterns, but it is still unknown whether they can be used to predict the distribution of soil organisms and functions across global biomes. Using a global field survey of 383 sites across contrasting climatic and vegetation conditions, here we showed that soil biodiversity and functions exhibited pervasive nonlinear patterns worldwide and are mainly governed by water availability (precipitation and potential evapotranspiration). Changes in water availability resulted in drastic shifts in soil biodiversity (bacteria, fungi, protists and invertebrates) and soil functions including plant-microbe interactions, plant productivity, soil biogeochemical cycles and soil carbon sequestration. Our findings highlight that crossing specific water availability thresholds can have critical consequences for the provision of essential ecosystem services needed to sustain our planet., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

Competing Interests: Conflict of interest The authors declare that they have no conflict of interest.

Knowing the extent and environmental drivers of forests is key to successfully restore degraded ecosystems, and to mitigate climate change and desertification impacts using tree planting. Water availability is the main limiting factor for the development of forests in drylands, yet the importance of groundwater resources and palaeoclimate as drivers of their current distribution has been neglected. Here we report that mid-Holocene climates and aquifer trends are key predictors of the distribution of dryland forests worldwide. We also updated the global extent of dryland forests to 1,283 million hectares and showed that failing to consider past climates and aquifers has resulted in ignoring or misplacing up to 130 million hectares of forests in drylands. Our findings highlight the importance of a wetter past and well-preserved aquifers to explain the current distribution of dryland forests, and can guide restoration actions by avoiding unsuitable areas for tree establishment in a drier world., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

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.)

Soil carbon losses to the atmosphere through soil respiration are expected to rise with ongoing temperature increases, but available evidence from mesic biomes suggests that such response disappears after a few years of experimental warming. However, there is lack of empirical basis for these temporal dynamics in soil respiration responses, and for the mechanisms underlying them, in drylands, which collectively form the largest biome on Earth and store 32% of the global soil organic carbon pool. We coupled data from a 10 year warming experiment in a biocrust-dominated dryland ecosystem with laboratory incubations to confront 0-2 years (short-term hereafter) versus 8-10 years (longer-term hereafter) soil respiration responses to warming. Our results showed that increased soil respiration rates with short-term warming observed in areas with high biocrust cover returned to control levels in the longer-term. Warming-induced increases in soil temperature were the main drivers of the short-term soil respiration responses, whereas longer-term soil respiration responses to warming were primarily driven by thermal acclimation and warming-induced reductions in biocrust cover. Our results highlight the importance of evaluating short- and longer-term soil respiration responses to warming as a mean to reduce the uncertainty in predicting the soil carbon-climate feedback in drylands., (© 2020 John Wiley & Sons Ltd.)

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.

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., (© 2019 John Wiley & Sons Ltd.)

Dispersal is a critical yet poorly understood factor underlying macroecological patterns in microbial communities 1 . Airborne microbial transport is assumed to occupy a central role in determining dispersal outcomes 2,3 , and extra-range dispersal has important implications for predicting ecosystem resilience and response to environmental change 4 . One of the most pertinent biomes in this regard is Antarctica, given its geographic isolation and vulnerability to climate change and human disturbance 5 . Here, we report microbial diversity in near-ground and high-altitude air above the largest ice-free Antarctic habitat, as well as that of underlying soil microbial communities. We found that persistent local airborne inputs were unable to fully explain Antarctic soil community assembly. Comparison with airborne microbial diversity from high-altitude and non-polar sources suggests that strong selection occurs during long-range atmospheric transport. The influence of selection during airborne transit and at sink locations varied between microbial phyla. Overall, the communities from this isolated Antarctic ecosystem displayed limited connectivity to the non-polar microbial pool, and alternative sources of recruitment are necessary to fully explain extant soil diversity. Our findings provide critical insights into the role of airborne transport limitation in determining microbial biogeographic patterns.

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., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

A positive soil carbon (C)-climate feedback is embedded into the climatic models of the IPCC. However, recent global syntheses indicate that the temperature sensitivity of soil respiration (R S ) in drylands, the largest biome on Earth, is actually lower in warmed than in control plots. Consequently, soil C losses with future warming are expected to be low compared with other biomes. Nevertheless, the empirical basis for these global extrapolations is still poor in drylands, due to the low number of field experiments testing the pathways behind the long-term responses of soil respiration (R S ) to warming. Importantly, global drylands are covered with biocrusts (communities formed by bryophytes, lichens, cyanobacteria, fungi, and bacteria), and thus, R S responses to warming may be driven by both autotrophic and heterotrophic pathways. Here, we evaluated the effects of 8-year experimental warming on R S , and the different pathways involved, in a biocrust-dominated dryland in southern Spain. We also assessed the overall impacts on soil organic C (SOC) accumulation over time. Across the years and biocrust cover levels, warming reduced R S by 0.30 μmol CO 2  m -2  s -1 (95% CI = -0.24 to 0.84), although the negative warming effects were only significant after 3 years of elevated temperatures in areas with low initial biocrust cover. We found support for different pathways regulating the warming-induced reduction in R S at areas with low (microbial thermal acclimation via reduced soil mass-specific respiration and β-glucosidase enzymatic activity) vs. high (microbial thermal acclimation jointly with a reduction in autotrophic respiration from decreased lichen cover) initial biocrust cover. Our 8-year experimental study shows a reduction in soil respiration with warming and highlights that biocrusts should be explicitly included in modeling efforts aimed to quantify the soil C-climate feedback in drylands., (© 2018 John Wiley & Sons Ltd.)

Drylands (hyperarid, arid, semiarid, and dry subhumid ecosystems) cover almost half of Earth's land surface and are highly vulnerable to environmental pressures. Here we provide an inventory of soil properties including carbon (C), nitrogen (N), and phosphorus (P) stocks within the current boundaries of drylands, aimed at serving as a benchmark in the face of future challenges including increased population, food security, desertification, and climate change. Aridity limits plant production and results in poorly developed soils, with coarse texture, low C:N and C:P, scarce organic matter, and high vulnerability to erosion. Dryland soils store 646 Pg of organic C to 2 m, the equivalent of 32% of the global soil organic C pool. The magnitude of the historic loss of C from dryland soils due to human land use and cover change and their typically low C:N and C:P suggest high potential to build up soil organic matter, but coarse soil textures may limit protection and stabilization processes. Restoring, preserving, and increasing soil organic matter in drylands may help slow down rising levels of atmospheric carbon dioxide by sequestering C, and is strongly needed to enhance food security and reduce the risk of land degradation and desertification.

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.

The relationship between the spatial variability of soil multifunctionality (i.e., the capacity of soils to conduct multiple functions; SVM) and major climatic drivers, such as temperature and aridity, has never been assessed globally in terrestrial ecosystems. We surveyed 236 dryland ecosystems from six continents to evaluate the relative importance of aridity and mean annual temperature, and of other abiotic (e.g., texture) and biotic (e.g., plant cover) variables as drivers of SVM, calculated as the averaged coefficient of variation for multiple soil variables linked to nutrient stocks and cycling. We found that increases in temperature and aridity were globally correlated to increases in SVM. Some of these climatic effects on SVM were direct, but others were indirectly driven through reductions in the number of vegetation patches and increases in soil sand content. The predictive capacity of our structural equation modelling was clearly higher for the spatial variability of N- than for C- and P-related soil variables. In the case of N cycling, the effects of temperature and aridity were both direct and indirect via changes in soil properties. For C and P, the effect of climate was mainly indirect via changes in plant attributes. These results suggest that future changes in climate may decouple the spatial availability of these elements for plants and microbes in dryland soils. Our findings significantly advance our understanding of the patterns and mechanisms driving SVM in drylands across the globe, which is critical for predicting changes in ecosystem functioning in response to climate change., (© 2018 by the Ecological Society of America.)

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.

The increase in aridity predicted with climate change will have a negative impact on the multiple functions and services (multifunctionality) provided by dryland ecosystems worldwide. In these ecosystems, soil communities dominated by mosses, lichens and cyanobacteria (biocrusts) play a key role in supporting multifunctionality. However, whether biocrusts can buffer the negative impacts of aridity on important biogeochemical processes controlling carbon (C), nitrogen (N), and phosphorus (P) pools and fluxes remains largely unknown. Here, we conducted an empirical study, using samples from three continents (North America, Europe and Australia), to evaluate how the increase in aridity predicted by climate change will alter the capacity of biocrust-forming mosses to modulate multiple ecosystem processes related to C, N and P cycles. Compared with soil surfaces lacking biocrusts, biocrust-forming mosses enhanced multiple functions related to C, N and P cycling and storage in semiarid and arid, but not in humid and dry-subhumid, environments. Most importantly, we found that the relative positive effects of biocrust-forming mosses on multifunctionality compared with bare soil increased with increasing aridity. These results were mediated by plant cover and the positive effects exerted by biocrust-forming mosses on the abundance of soil bacteria and fungi. Our findings provide strong evidence that the maintenance of biocrusts is crucial to buffer negative effects of climate change on multifunctionality in global drylands., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

Soil bacteria and fungi play key roles in the functioning of terrestrial ecosystems, yet our understanding of their responses to climate change lags significantly behind that of other organisms. This gap in our understanding is particularly true for drylands, which occupy ∼41% of Earth´s surface, because no global, systematic assessments of the joint diversity of soil bacteria and fungi have been conducted in these environments to date. Here we present results from a study conducted across 80 dryland sites from all continents, except Antarctica, to assess how changes in aridity affect the composition, abundance, and diversity of soil bacteria and fungi. The diversity and abundance of soil bacteria and fungi was reduced as aridity increased. These results were largely driven by the negative impacts of aridity on soil organic carbon content, which positively affected the abundance and diversity of both bacteria and fungi. Aridity promoted shifts in the composition of soil bacteria, with increases in the relative abundance of Chloroflexi and α-Proteobacteria and decreases in Acidobacteria and Verrucomicrobia. Contrary to what has been reported by previous continental and global-scale studies, soil pH was not a major driver of bacterial diversity, and fungal communities were dominated by Ascomycota. Our results fill a critical gap in our understanding of soil microbial communities in terrestrial ecosystems. They suggest that changes in aridity, such as those predicted by climate-change models, may reduce microbial abundance and diversity, a response that will likely impact the provision of key ecosystem services by global drylands.

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.)

Nitrogen (N) deposition has doubled the natural N inputs received by ecosystems through biological N fixation and is currently a global problem that is affecting the Mediterranean regions. We evaluated the existing relationships between increased atmospheric N deposition and biogeochemical indicators related to soil chemical factors and cryptogam species across semiarid central, southern, and eastern Spain. The cryptogam species studied were the biocrust-forming species Pleurochaete squarrosa (moss) and Cladonia foliacea (lichen). Sampling sites were chosen in Quercus coccifera (kermes oak) shrublands and Pinus halepensis (Aleppo pine) forests to cover a range of inorganic N deposition representative of the levels found in the Iberian Peninsula (between 4.4 and 8.1 kg N ha(-1) year(-1)). We extended the ambient N deposition gradient by including experimental plots to which N had been added for 3 years at rates of 10, 20, and 50 kg N ha(-1) year(-1). Overall, N deposition (extant plus simulated) increased soil inorganic N availability and caused soil acidification. Nitrogen deposition increased phosphomonoesterase (PME) enzyme activity and PME/nitrate reductase (NR) ratio in both species, whereas the NR activity was reduced only in the moss. Responses of PME and NR activities were attributed to an induced N to phosphorus imbalance and to N saturation, respectively. When only considering the ambient N deposition, soil organic C and N contents were positively related to N deposition, a response driven by pine forests. The PME/NR ratios of the moss were better predictors of N deposition rates than PME or NR activities alone in shrublands, whereas no correlation between N deposition and the lichen physiology was observed. We conclude that integrative physiological measurements, such as PME/NR ratios, measured on sensitive species such as P. squarrosa, can provide useful data for national-scale biomonitoring programs, whereas soil acidification and soil C and N storage could be useful as additional corroborating ecosystem indicators of chronic N pollution.

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.)

The biogeochemical cycles of carbon (C), nitrogen (N) and phosphorus (P) are interlinked by primary production, respiration and decomposition in terrestrial ecosystems. It has been suggested that the C, N and P cycles could become uncoupled under rapid climate change because of the different degrees of control exerted on the supply of these elements by biological and geochemical processes. Climatic controls on biogeochemical cycles are particularly relevant in arid, semi-arid and dry sub-humid ecosystems (drylands) because their biological activity is mainly driven by water availability. The increase in aridity predicted for the twenty-first century in many drylands worldwide may therefore threaten the balance between these cycles, differentially affecting the availability of essential nutrients. Here we evaluate how aridity affects the balance between C, N and P in soils collected from 224 dryland sites from all continents except Antarctica. We find a negative effect of aridity on the concentration of soil organic C and total N, but a positive effect on the concentration of inorganic P. Aridity is negatively related to plant cover, which may favour the dominance of physical processes such as rock weathering, a major source of P to ecosystems, over biological processes that provide more C and N, such as litter decomposition. Our findings suggest that any predicted increase in aridity with climate change will probably reduce the concentrations of N and C in global drylands, but increase that of P. These changes would uncouple the C, N and P cycles in drylands and could negatively affect the provision of key services provided by these ecosystems.

An integrated perspective of the most important factors driving the abundance of ammonia-oxidizing bacteria (AOB) and archaea (AOA) in natural ecosystems is lacking, especially in drylands. We evaluated how different climatic, abiotic, and nutrient-related factors determine AOA and AOB abundance in bare and vegetated microsites from grasslands throughout the Mediterranean Basin. We found a strong negative relationship between the abundance of AOA genes and soil fertility (availability of C, N, and P). Aridity and other abiotic factors (pH, sand content, and electrical conductivity) were more important than soil fertility in modulating the AOA/AOB ratio. AOB were more abundant under vegetated microsites, while AOA, highly resistant to stressful conditions, were more abundant in bare ground areas. These results suggest that AOA may carry out nitrification in less fertile microsites, while AOB predominate under more fertile conditions. Our results indicate that the influence of aridity and pH on the relative dominance of AOA and AOB genes is ultimately determined by local-scale environmental changes promoted by perennial vegetation. Thus, in spatially heterogeneous ecosystems such as drylands, there is a mutual exclusion and niche division between these microorganisms, suggesting that they may be functionally complementary., (© 2013 Federation of European Microbiological Societies. Published by John Wiley & Sons Ltd. All rights reserved.)

Biological soil crusts (BSCs) are key biotic components of dryland ecosystems worldwide that control many functional processes, including carbon and nitrogen cycling, soil stabilization and infiltration. Regardless of their ecological importance and prevalence in drylands, very few studies have explicitly evaluated how climate change will affect the structure and composition of BSCs, and the functioning of their constituents. Using a manipulative experiment conducted over 3 years in a semi-arid site from central Spain, we evaluated how the composition, structure and performance of lichen-dominated BSCs respond to a 2.4°C increase in temperature, and to an approximately 30 per cent reduction of total annual rainfall. In areas with well-developed BSCs, warming promoted a significant decrease in the richness and diversity of the whole BSC community. This was accompanied by important compositional changes, as the cover of lichens suffered a substantial decrease with warming (from 70 to 40% on average), while that of mosses increased slightly (from 0.3 to 7% on average). The physiological performance of the BSC community, evaluated using chlorophyll fluorescence, increased with warming during the first year of the experiment, but did not respond to rainfall reduction. Our results indicate that ongoing climate change will strongly affect the diversity and composition of BSC communities, as well as their recovery after disturbances. The expected changes in richness and composition under warming could reduce or even reverse the positive effects of BSCs on important soil processes. Thus, these changes are likely to promote an overall reduction in ecosystem processes that sustain and control nutrient cycling, soil stabilization and water dynamics.

Species interactions play key roles in linking the responses of populations, communities, and ecosystems to environmental change. For instance, species interactions are an important determinant of the complexity of changes in trophic biomass with variation in resources. Water resources are a major driver of terrestrial ecology and climate change is expected to greatly alter the distribution of this critical resource. While previous studies have documented strong effects of global environmental change on species interactions in general, responses can vary from region to region. Dryland ecosystems occupy more than one-third of the Earth's land mass, are greatly affected by changes in water availability, and are predicted to be hotspots of climate change. Thus, it is imperative to understand the effects of environmental change on these globally significant ecosystems. Here, we review studies of the responses of population-level plant-plant, plant-herbivore, and predator-prey interactions to changes in water availability in dryland environments in order to develop new hypotheses and predictions to guide future research. To help explain patterns of interaction outcomes, we developed a conceptual model that views interaction outcomes as shifting between (1) competition and facilitation (plant-plant), (2) herbivory, neutralism, or mutualism (plant-herbivore), or (3) neutralism and predation (predator-prey), as water availability crosses physiological, behavioural, or population-density thresholds. We link our conceptual model to hypothetical scenarios of current and future water availability to make testable predictions about the influence of changes in water availability on species interactions. We also examine potential implications of our conceptual model for the relative importance of top-down effects and the linearity of patterns of change in trophic biomass with changes in water availability. Finally, we highlight key research needs and some possible broader impacts of our findings. Overall, we hope to stimulate and guide future research that links changes in water availability to patterns of species interactions and the dynamics of populations and communities in dryland ecosystems., (© 2011 The Authors. Biological Reviews © 2011 Cambridge Philosophical Society.)

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.

Roadside grasslands undergoing secondary succession are abundant, and represent ecologically meaningful examples of novel, human-created ecosystems. Interactions between plant and soil communities (hereafter plant-soil interactions) are of major importance in understanding the role of biotic control in ecosystem functioning, but little is known about these links in the context of ecosystem restoration and succession. The assessment of the key biotic communities and interactions driving ecosystem development will help practitioners to better allocate the limited resources devoted to roadside grassland restoration. We surveyed roadside grasslands from three successional stages (0-2, 7-9, and >20 years) in two Mediterranean regions of Spain. Structural equation modeling was used to evaluate how interactions between plants, biological soil crusts (BSCs), and soil microbial functional diversity (soil microorganisms) affect indicators of ecosystem development and restoration: plant similarity to the reference ecosystem, erosion control, and soil C storage and N accumulation. Changes in plant community composition along the successional gradient exerted the strongest influence on these indicators. High BSC cover was associated with high soil stability, and high soil microbial functional diversity from late-successional stages was associated with high soil fertility. Contrary to our expectations, the indirect effects of plants, mediated by either BSCs or soil microorganisms, were very weak in both regions, suggesting a minor role for plant-soil interactions upon ecosystem development indicators over long periods. Our results suggest that natural vegetation dynamics effectively improved ecosystem development within a time frame of 20 years in the grasslands evaluated. They also indicate that this time could be shortened if management actions focus on: (1) maintaining well-conserved natural areas close to roadsides to enhance plant compositional changes towards late-successional stages, (2) increasing BSC cover in areas under strong erosion risk, to avoid soil loss, and (3) enhancing soil microbial functional diversity in resource-limited areas, to enhance soil C and N accumulation.

Encroachment of woody plants into grasslands has generated considerable interest among ecologists. Syntheses of encroachment effects on ecosystem processes have been limited in extent and confined largely to pastoral land uses or particular geographical regions. We used univariate analyses, meta-analysis and structural equation modelling to test the propositions that (1) shrub encroachment does not necessarily lead to declines in ecosystem functions and (2) shrub traits influence the functional outcome of encroachment. Analyses of 43 ecosystem attributes from 244 case studies worldwide showed that some attributes consistently increased with encroachment (e.g. soil C, N), and others declined (e.g. grass cover, pH), but most exhibited variable responses. Traits of shrubs were associated with significant, though weak, structural and functional outcomes of encroachment. Our review revealed that encroachment had mixed effects on ecosystem structure and functioning at global scales, and that shrub traits influence the functional outcome of encroachment. Thus, a simple designation of encroachment as a process leading to functionally, structurally or contextually degraded ecosystems is not supported by a critical analysis of existing literature. Our results highlight that the commonly established link between shrub encroachment and degradation is not universal., (© 2011 Blackwell Publishing Ltd/CNRS and State Government of NSW.)

Background and Aims: Competition drives self-thinning (density-dependent mortality) in crowded plant populations. Facilitative interactions have been shown to affect many processes in plant populations and communities, but their effects on self-thinning trajectories have not been investigated., Methods: Using an individual-based 'zone-of-influence' model, we studied the potential effects of the size symmetry of competition, abiotic stress and facilitation on self-thinning trajectories in plant monocultures. In the model, abiotic stress reduced the growth of all individuals and facilitation ameliorated the effects of stress on interacting individuals., Key Results: Abiotic stress made the log biomass-log density relationship during self-thinning steeper, but this effect was reduced by positive interactions among individuals. Size-asymmetric competition also influenced the self-thinning slope., Conclusions: Although competition drives self-thinning, its course can be affected by abiotic stress, facilitation and competitive symmetry.

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.

The worldwide phenomenon of shrub encroachment in grass-dominated dryland ecosystems is commonly associated with desertification. Studies of the purported desertification effects associated with shrub encroachment are often restricted to relatively few study areas, and document a narrow range of possible impacts upon biota and ecosystem processes. We conducted a study in degraded Mediterranean grasslands dominated by Stipa tenacissima to simultaneously evaluate the effects of shrub encroachment on the structure and composition of multiple biotic community components, and on various indicators of ecosystem function. Shrub encroachment enhanced vascular plant richness, biomass of fungi, actinomycetes and other bacteria, and was linked with greater soil fertility and N mineralization rates. While shrub encroachment may be a widespread phenomenon in drylands, an interpretation that this is an expression of desertification is not universal. Our results suggest that shrub establishment may be an important step in the reversal of desertification processes in the Mediterranean region.

The monitoring of desertification processes, and particularly the development of "early-warning" systems, is an increasingly important development in the management of drylands. It has been shown that the patch size distribution of dryland vegetation can be described using power laws and that deviations from such patterns may be used as an early-warning signal for the onset of desertification. We tested this idea using data from 29 semiarid steppes located along a latitudinal gradient in Spain. A truncated power law (TPL) fitted the patch size distribution of perennial vegetation better than a power law in all the evaluated sites. Variations in this distribution, as measured with the scaling exponent (gamma) of the TPL, were not related to total perennial cover, but a negative logarithmic relationship was found between gamma and soil variables related to desertification processes (total nitrogen, total phosphorus, and organic carbon). A positive and stronger linear relationship was found between total perennial cover and the same soil variables. Our results suggest that deviations from a patch size distribution characterized by a power law are not directly related to desertification. They also indicate that plant cover can be used to effectively monitor key variables linked to desertification processes.

We introduce a spatially explicit model that evaluates how the trade-offs between the life strategies of two interacting plant species affect the outcome of their interaction along environmental severity gradients. In our model, we represent the landscape as a two-dimensional lattice, with environmental severity increasing from left to right. Two species with different strategies, a competitor and a stress-tolerant, interact in the lattice. We find that facilitation expands the realized niche of the competitor into harsh environments by suppressing the stress-tolerant species. Most of their coexisting range is dominated by a positive effect of one species on another, with a reciprocal negative effect from the species receiving the benefits on its benefactor ("+, -"), whereas mutualistic ("+, +") interactions are only found in the harshest part of the environmental gradient. Contrarily as assumed by models commonly used in facilitation research (e.g. dual-lattice models), our results indicate that "+, +" interactions are not dominant, and that their differences with "+, -" interactions along environmental severity gradients depend on the strategies of the interacting species. By integrating the trade-off between competitive ability and stress tolerance, our model provides a new framework to investigate the interplay of facilitative and competitive interactions along environmental gradients and their impacts on processes such as population dynamics and community organization.

Theories based on competition for resources predict a monotonic negative relationship between population density and individual biomass in plant populations. They do not consider the role of facilitative interactions, which are known to be important in high stress environments. Using an individual-based 'zone-of-influence' model, we investigated the hypothesis that the balance between facilitative and competitive interactions determines biomass-density relationships. We tested model predictions with a field experiment on the clonal grass Elymus nutans in an alpine meadow. In the model, the relationship between mean individual biomass and density shifted from monotonic to humped as abiotic stress increased. The model results were supported by the field experiment, in which the greatest individual and population biomass were found at intermediate densities in a high-stress alpine habitat. Our results show that facilitation can affect biomass-density relationships.

Background and Aims: Neutral theory predicts that the diversity and relative abundance of species in ecological communities do not depend on their specific traits. This prediction remains controversial, as many studies suggest that variations in the niches of species determine the structure of communities. The aim of this study was to test empirically the relative importance of niche and neutral processes as drivers of species abundance within plant communities along a successional gradient., Methods: Information on the abundance (density and frequency) and traits (aboveground individual biomass and seed mass) of > 90 species was collected in alpine and sub-alpine meadows of the Tibet Plateau (China). A successional gradient (1, 3, 15 and 30 years after abandonment) was established in a sub-alpine meadow. The relationships between species traits and their abundance were evaluated using regression models., Key Results: Seed mass was negatively related to both species density (r = -0.6270, P < 0.001) and frequency (r = -0.5335, P = 0.005) in the 1-year meadow. Such relationships disappeared along the successional gradient evaluated (P > 0.07 in the 3-, 15- and 30-year meadows). Data gathered in all sites showed a significant negative relationship between the average individual biomass of a given species and its density within the community (r < -0.30, P < 0.025 in all cases)., Conclusions: The results show that seed mass was a key driver of species abundance in early successional communities, and that niche forces may become more important as succession progresses. They also indicate that predictions from neutral theory, in its current form, do not hold for the meadow communities studied.

In this millennium, global drylands face a myriad of problems that present tough research, management, and policy challenges. Recent advances in dryland development, however, together with the integrative approaches of global change and sustainability science, suggest that concerns about land degradation, poverty, safeguarding biodiversity, and protecting the culture of 2.5 billion people can be confronted with renewed optimism. We review recent lessons about the functioning of dryland ecosystems and the livelihood systems of their human residents and introduce a new synthetic framework, the Drylands Development Paradigm (DDP). The DDP, supported by a growing and well-documented set of tools for policy and management action, helps navigate the inherent complexity of desertification and dryland development, identifying and synthesizing those factors important to research, management, and policy communities.

While it is well-established that the spatial distribution of soil nutrients (soil heterogeneity) influences the competitive ability and survival of individual plants, as well as the productivity of plant communities, there is a paucity of data on how soil heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. To evaluate the effects of elevated CO(2), soil heterogeneity and diversity (species richness and composition) on productivity, patterns of biomass allocation and root foraging precision, we conducted an experiment with grassland assemblages formed by monocultures, two- and three-species mixtures of Lolium perenne, Plantago lanceolata and Holcus lanatus. The experiment lasted for 90 days, and was conducted on microcosms built out of PVC pipe (length 38 cm, internal diameter 10 cm). When nutrients were heterogeneously supplied (in discrete patches), assemblages exhibited precise root foraging patterns, and had higher total, above- and belowground biomass. Greater aboveground biomass was observed under elevated CO(2). Species composition affected the below:aboveground biomass ratio and interacted with nutrient heterogeneity to determine belowground and total biomass. Species richness had no significant effects, and did not interact with either CO(2) or nutrient heterogeneity. Under elevated CO(2) conditions, the two- and three-species mixtures showed a clear trend towards underyielding. Our results show that differences among composition levels were dependent on soil heterogeneity, highlighting its potential role in modulating diversity-productivity relationships.

Patterns of resource availability and heterogeneity shape the composition, productivity, and dynamics of plant assemblages in a wide variety of terrestrial ecosystems. Despite this, the responses of plant assemblages to simultaneous changes in the availability and heterogeneity of more than a single resource are virtually unknown. To fill this gap, microcosms consisting of assemblages formed by Lolium perenne, Plantago lanceolata, Anthoxantum odoratum, Holcus lanatus, and Trifolium repens were grown in a factorial experiment with the following treatments: nutrient availability (NA), water availability (WA), spatial nutrient heterogeneity (NH), and temporal water heterogeneity (WH). Assemblages exhibited precise root foraging patterns in response to nutrient heterogeneity, which were modified by NA and WA. A series of two- and three-way interactions involving the four factors evaluated determined biomass production, the belowground: aboveground biomass ratio, the patterns of root biomass allocation with depth, and the relative contribution to aboveground biomass of Lolium and Anthoxanthum. In all cases, these interactions explained significant amounts of the variation found in the data. Our study demonstrates that considering the interactions between resource availability and heterogeneity allows for a refinement of predictions that can detectably reduce the error associated with extrapolating from single factor analyses.

Biological soil crusts are very sensitive to human-induced disturbances and are in a degraded state in many areas throughout their range. Given their importance in the functioning of arid and semiarid ecosystems, restoring these crusts may contribute to the recovery of ecosystem functionality in degraded areas. We conducted a factorial microcosm experiment to evaluate the effects of inoculation type (discrete fragments vs slurry), fertilization (control vs addition of composted sewage sludge), and watering frequency (two vs five times per week) on the cyanobacterial composition, nitrogen fixation, chlorophyll content, and net CO2 exchange rate of biological soil crusts inoculated on a semiarid degraded soil from SE Spain. Six months after the inoculation, the highest rates of nitrogen fixation and chlorophyll a content were found when the biological crusts were inoculated as slurry, composted sewage sludge was added, and the microcosms were watered five times per week. Net CO2 exchange rate increased when biological crusts were inoculated as slurry and the microcosms were watered five times per week. Denaturing gradient gel electrophoresis fingerprints and phylogenetic analyses indicated that most of the cyanobacterial species already present in the inoculated crust had the capability to spread and colonize the surface of the surrounding soil. These analyses showed that cyanobacterial communities were less diverse when the microcosms were watered five times per week, and that watering frequency (followed in importance by the addition of composted sewage sludge and inoculation type) was the treatment that most strongly influenced their composition. Our results suggest that the inoculation of biological soil crusts in the form of slurry combined with the addition of composted sewage sludge could be a suitable technique to accelerate the recovery of the composition and functioning of biological soil crusts in drylands.

Background and Aims: Size-asymmetric competition occurs when larger plants have a disproportionate advantage in competition with smaller plants. It has been hypothesized that nutrient heterogeneity may promote it. Experiments testing this hypothesis are inconclusive, and in most cases have evaluated the effects of nutrient heterogeneity separately from other environmental factors. The aim of this study was to test, using populations of Lolium perenne, Plantago lanceolata and Holcus lanatus, two hypotheses: (a) nutrient heterogeneity promotes size-asymmetric competition; and (b) nutrient heterogeneity interacts with both atmospheric CO2 partial pressure (P(CO2)) and nutrient availability to determine the magnitude of this response., Methods: Microcosms consisting of monocultures of the three species were grown for 90 d in a factorial experiment with the following treatments: P(CO2) (37.5 and 70 Pa) and nutrient availability (NA; 40 and 120 mg of N added as organic material) combined with different spatial distribution of the organic material (NH; homogeneous and heterogeneous). Differences in the size of individual plants within populations (size inequality) were quantified using the coefficient of variation of individual above-ground biomass and the combined biomass of the two largest individuals in each microcosm. Increases in size inequality were associated with size-asymmetric competition., Key Results: Size inequality increased when the nutrients were heterogeneously supplied in the three species. The effects of NH on this response were more pronounced under high nutrient supply in both Plantago and Holcus (significant NA x NH interactions) and under elevated P(CO2) in Plantago (significant P(CO2) x NA x NH interaction). No significant two- and three-way interactions were found for Lolium., Conclusions: Our first hypothesis was supported by our results, as nutrient heterogeneity promoted size-asymmetric competition in the three species evaluated. Nutrient supply and P(CO2) modified the magnitude of this effect in Plantago and Holcus, but not in Lolium. Thus, our second hypothesis was partially supported.

Theoretical models predict that the relative importance of facilitation and competition may vary inversely across gradients of abiotic stress. However, these predictions have not been thoroughly tested in the field, especially in semi-arid environments. In this study, we evaluated how the net effect of the tussock grass Stipa tenacissima on the shrub Pistacia lentiscus varied across a gradient of abiotic stress in semi-arid Mediterranean steppes. We fitted the relationship between accumulated rainfall and the relative neighbour index (our measures of abiotic stress and of the net effect of S. tenacissima on P. lentiscus, respectively), which varied across this gradient, to a quadratic model. Competitive interactions dominated at both extremes of the gradient. Our results do not support established theory. Instead, they suggest that a shift from facilitation to competition under high abiotic stress conditions is likely to occur when the levels of the most limiting resource are so low that the benefits provided by the facilitator cannot overcome its own resource uptake.

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.