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.

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 climatechange models, may reduce microbial abundance and diversity, a response that will likely impact the provision of key ecosystem services by global drylands.bacteria | fungi | climate change | arid | semiarid

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.

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Abstract. The purpose of this study was to relate regional variation in litter mass-loss rates (first year) in pine forests to climate across a large, continental-scale area. The variation in mass-loss rate was analyzed using 39 experimental sites spanning climatic regions from the subarctic to subtropical and Mediterranean: the latitudinal gradient ranged from 31 N to 70 N and may represent the the largest geographical area that has ever been sampled and observed for the purpose of studying biogeochemical processes. Because of unified site design and uniform laboratory procedures, data from all sites were directly comparable and permitted a determination of the relative influence of climate versus substrate quality viewed from the perspective of broad regional scales.Simple correlation applied to the entire data set indicated that annual actual evapotranspiration (AET) should be the leading climatic constraint on mass-loss rates (R2d = 128 0.496). The combination of AET, average July temp. and average annual temp. could explain about 70% of the sites' variability on litter mass-loss. In an analysis of 23 Scots pine sites north of the Alps and Carpatians AET alone could account for about 65% of the variation and the addition of a substrate-quality variable was sufficiently significant to be used in a model.The influence of litter quality was introduced into a model, using data from 11 sites at which litter of different quality had been incubated. These sites are found in Germany, the Netherlands, Sweden and Finland. At any one site most (> 90%/6) of the variation in mass-loss rates could be explained by one of the litter-quality variables giving concentration of nitrogen, phosphorus or water solubles. However, even when these models included nitrogen or phosphorus even small changes in potential evapotranspiration resulted in large changes in early-phase decay rates.Further regional subdivision of the data set, resulted in a range of strength in the relationship between loss rate and climatic variables, from very weak in Central Europe to strong for the Scandinavian and Atlantic coast sites (Rdj = 0.912; AET versus litter mass loss). Much of the variation in observed loss rates could be related to continental versus marine/Atlantic influences. Inland locations had mass-loss rates lower than should be expected on the basis of for example AET alone. Attempts to include seasonality variables were not successful. It is clear that either unknown errors and biases, or, unknown variables are causing these regional differences in response to climatic variables. Nevertheless these results show the powerful influence of climate as a control of the broad-scale geography of mass-loss rates and substrate quality at the stand level.Some of these relationships between mass-loss rate and climatic variables are among the highest ever reported, probably because of the care taken to select uniform sites and experimental methods. This suggest that superior, base line maps of predicted mass-loss rates could be produced using climatic...

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.

Understanding how drylands respond to ongoing environmental change is extremely important for global sustainability. Here we review 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, illustrate how N deposition and grazing pressure are impacting ecosystem functioning in drylands worldwide, and highlight 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.

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 realworld ecosystems remains unclear. We used two large datasets 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 * corresponding author: santiago.soliveres@ips.unibe.ch. Authorship SS, EA and FTM conceived the study, SS and WU analyzed the data, SS, MF, FTM, DP, MB, SB, JLQ, IS, MDB, WW, VO, PM, SAS and JM gathered data, SS wrote the first draft and all co-authors significantly contributed to improve it. Europe PMC Funders Group Europe PMC Funders Author ManuscriptsEurope PMC Funders Author Manuscripts 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 homogenization.