Drivers of non-native plant success in drylands are poorly understood. Here we identify functional differences between dryland native and non-native perennial plants and assess how biotic, abiotic and anthropogenic factors shape the success of the latter. On the basis of plant community and functional trait data from 98 sites across 25 countries, we report a total of 41 non-native plant species at 31 sites. Non-natives tend towards faster growth strategies than natives. Non-native plant richness is higher at sites with greater grazing pressure and under environmental conditions associated with higher soil fertility, decomposition and fungal richness-conditions that tend to occur in less arid regions-and lower where native plant and herbivore richness are greater. Non-native plant cover correlates positively with grazing pressure and negatively with native plant richness. Taken together, our results suggest that non-native plant success in drylands is facilitated when high grazing pressure coincides with elevated resource availability. Such context-dependence of non-native plant success and linkages with native plant and herbivore diversity highlight the need for managing grazing and conserving biodiversity across the world's drylands.

Biocrusts cover a substantial proportion of dryland soil surfaces, but the relationships between these surface communities and soil salinity and alkalinity remain poorly understood. In this study, we used high-throughput DNA sequencing to investigate the bacterial communities within biocrusts from saline-alkaline soils surrounding playas in northeast Spain (Alcañiz region). Our aims were to examine whether these soils support the development of specific biocrust bacterial communities, and how these communities relate to biocrust types (cyanobacterial, lichen, and moss-dominated) and their ecological functions. The results showed that biocrusts in these saline-alkaline soils harboured bacterial communities characterized by low cyanobacterial relative abundances (0.5–5.0%), and were composed of many rare species (311–385 OTUs) and only a few dominant species (16–18 OTUs), while habitat generalists (50 OTUs) and specialists (8–20 OTUs) were detected both across the rare and dominant species. Community assembly appeared to be shaped by deterministic environmental filtering and a secondary contribution from stochastic processes (stochasticity ratio 0.28–0.36), and the compositionally-differentiated bacterial communities among biocrust types suggest a potential specific link between bacterial assemblages and biocrust photoautotrophs. Furthermore, a clear co-variation between biocrust development and soil salinity and alkalinity supports a shift in community composition from more specialized to more generalized taxa along a decreasing salinity and alkalinity gradient. Compared with cyanobacterial soil crusts, isotope measurements revealed higher δ13C but lower δ15N in lichen and/or moss soil crusts (P < 0.05), with organic carbon and total nitrogen higher but salinity and alkalinity lower (P < 0.05). Collectively, our results highlight that diverse biocrust types can develop on saline-alkaline soils, and that specialized bacterial community composition and function are associated with biocrust types as well as soil salinity and/or alkalinity. These findings suggest specific bacterial salinity and/or alkalinity preferences may assist in selecting appropriate microbial approaches for biocrust restoration and soil functional management in these and other saline-alkaline environments.

Mineral-associated organic carbon (MAOC) constitutes a major fraction of global soil carbon and is assumed less sensitive to climate than particulate organic carbon (POC) due to protection by minerals. Despite its importance for long-term carbon storage, the response of MAOC to changing climates in drylands, which cover more than 40% of the global land area, remains unexplored. Here we assess topsoil organic carbon fractions across global drylands using a standardized field survey in 326 plots from 25 countries and 6 continents. We find that soil biogeochemistry explained the majority of variation in both MAOC and POC. Both carbon fractions decreased with increases in mean annual temperature and reductions in precipitation, with MAOC responding similarly to POC. Therefore, our results suggest that ongoing climate warming and aridification may result in unforeseen carbon losses across global drylands, and that the protective role of minerals may not dampen these effects..

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.

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

Increases in the abundance of woody species have been reported to affect the provisioning of ecosystem services in drylands worldwide. However, it is virtually unknown how multiple biotic and abiotic drivers, such as climate, grazing, and fire, interact to determine woody dominance across global drylands. We conducted a standardized field survey in 304 plots across 25 countries to assess how climatic features, soil properties, grazing, and fire affect woody dominance in dryland rangelands. Precipitation, temperature, and grazing were key determinants of tree and shrub dominance. The effects of grazing were determined not solely by grazing pressure but also by the dominant livestock species. Interactions between soil, climate, and grazing and differences in responses to these factors between trees and shrubs were key to understanding changes in woody dominance. Our findings suggest that projected changes in climate and grazing pressure may increase woody dominance in drylands, altering their structure and functioning.

Biological soil crusts, or biocrusts, are microbial communities found in soil surfaces in drylands and in other locations where vascular plant cover is incomplete. They are functionally significant for numerous ecosystem services, most notably in the C fixation and storage due to the ubiquity of photosynthetic microbes. Whereas carbon fixation and storage have been well studied in biocrusts, the composition, function and characteristics of other organisms in the biocrust such as heterotrophic bacteria and especially fungi are considerably less studied and this limits our ability to gain a holistic understanding of biocrust ecology and function. In this research we characterised the fungal community in biocrusts developed on Kalahari Sand soils from a site in southwest Botswana, and combined these data with previously published bacterial community data from the same site. By identifying organisational patterns in the community structure of fungi and bacteria, we found fungi that were either significantly associated with biocrust or the soil beneath biocrusts, leading to the conclusion that they likely perform functions related to the spatial organisation observed. Furthermore, we showed that within biocrusts bacterial and fungal community structures are correlated with each other i.e., a change in the bacterial community is reflected by a corresponding change in the fungal community. Importantly, this correlation but that this correlation does not occur in nearby soils. We propose that different fungi engage in short-range and long-range interactions with dryland soil surface bacteria. We have identified fungi which are candidates for further studies into their potential roles in biocrust ecology at short ranges (e.g., processing of complex compounds for waste management and resource provisioning) and longer ranges (e.g., translocation of resources such as water and the fungal loop model). This research shows that fungi are likely to have a greater contribution to biocrust function and dryland ecology than has generally been recognised.

Research in global change ecology relies heavily on global climatic grids derived from estimates of air temperature in open areas at around 2 m above the ground. These climatic grids do not reflect conditions below vegetation canopies and near the ground surface, where critical ecosystem functions occur and most terrestrial species reside. Here, we provide global maps of soil temperature and bioclimatic variables at a 1‐km 2 resolution for 0–5 and 5–15 cm soil depth. These maps were created by calculating the difference (i.e. offset) between in situ soil temperature measurements, based on time series from over 1200 1‐km 2 pixels (summarized from 8519 unique temperature sensors) across all the world's major terrestrial biomes, and coarse‐grained air temperature estimates from ERA5‐Land (an atmospheric reanalysis by the European Centre for Medium‐Range Weather Forecasts). We show that mean annual soil temperature differs markedly from the corresponding gridded air temperature, by up to 10°C (mean = 3.0 ± 2.1°C), with substantial variation across biomes and seasons. Over the year, soils in cold and/or dry biomes are substantially warmer (+3.6 ± 2.3°C) than gridded air temperature, whereas soils in warm and humid environments are on average slightly cooler (−0.7 ± 2.3°C). The observed substantial and biome‐specific offsets emphasize that the projected impacts of climate and climate change on near‐surface biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments. The global soil‐related bioclimatic variables provided here are an important step forward for any application in ecology and related disciplines. Nevertheless, we highlight the need to fill remaining geographic gaps by collecting more in situ measurements of microclimate conditions to further enhance the spatiotemporal resolution of global soil temperature products for ecological applications.

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 speciespoor 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.

Soil biocrusts, formed from communities of microbes and their extracellular products are a common feature of dryland soil surfaces. Biocrust organisms are only intermittently metabolically active, but due to their ubiquity they make a significant contribution to the carbon cycle. Quantification of the controls and insights into the interlinked process of photosynthesis and respiration are essential to enhancing our understanding of the carbon cycle in the world's drylands. Yet, there have been relatively few field studies investigating controls on both biocrust photosynthesis and respiration. We undertook field-based experiments at two dune sites during the dry season in Diamantina National Park in Queensland, Australia to determine how biocrust hydration and illumination affect soil CO2 flux and photosynthesis. Static chambers and an infra-red gas analyser were used to quantify soil CO2 flux, and a fluorometer and a CFlmager were used to determine a range of photosynthetic parameters in the field and laboratory respectively. When dry, biocrust photosynthetic activity was not detected and soil CO2 flux was very low irrespective of biocrust cover. Hydration led to a large and immediate increase in CO2 flux, which was more pronounced in the presence of biocrusts and on the dune with thinner biocrusts. Hydration also initiated the onset of photosynthesis in some biocrusts, which was greatest under low light conditions and sustained with further hydration. There were only infrequent periods of net CO2 uptake to the soil, occurring when CO2 uptake due to photosynthetic activity was less than background soil CO2 flux. Chlorophyll fluorescence imaging indicated biocrust spatial heterogeneity was evident at the cm scale where microtopography creates a myriad of environments for different crust organisms. Our findings demonstrate that biocrusts are highly spatially heterogenetic at both landscape and small scale, which suggests the maintenance of biocrust spatial diversity is likely to be key to imparting resilience to changing climate and disturbance. As well as reaffirming the importance of biocrusts for the carbon cycle in dryland dune soils the study demonstrates that biocrust respiration and photosynthesis respond differently to hydration and shading. This adds an unpredictability to the distribution of soil carbon stocks and the gaseous exchanges of CO2 between the surface and atmosphere. Future changes to precipitation and increased temperatures are likely to reduce soil moisture across much of the Australian interior and consequently biocrusts may experience a decline in biomass, structure, and function which could have significant repercussions beyond carbon stocks.

Numerous permanent and temporary wetlands occur throughout the world’s drylands. Although characterised by diverse water inundation frequencies, durations and depths, these wetlands in drylands are typically hotspots of biological activity and productivity. The healthy functioning and even existence of many wetlands in drylands, however, is threatened by desiccation resulting from a combination of climate change and human disturbance. Near Alcañiz in arid northeast Spain, three adjacent saladas (playas) with contrasting hydroperiods provide an opportunity to investigate how moisture availability affects their soil carbon (C) stocks, CO2 efflux, and microbial communities. Predicting the impacts of changes in moisture availability on the C cycle in wetlands in drylands is challenging but important because many contain large C stocks and may be significant sources and sinks of greenhouse gases. Frequent inundation and/or near-permanent soil water saturation supports the generation of organic C from a range of different sources. Soil inorganic C was greatest on the driest salada (3.8%) compared to the wetter saladas (3.0% and 2.1%) owing to evaporative concentration and the reaction of CO2 with available Ca2+, Mg2+ and Na+ ions. CO2 efflux was greatest at intermediate moisture levels (142 mg CO2 m−2 h−1), but the spatial and temporal variability in CO2 efflux on salada surfaces is large, demonstrating the need for intensive sampling regimes to provide realistic estimates of their contribution to atmospheric CO2 exchanges. Different microbial community structures also characterise each salada. The saladas near Alcañiz, and many other similar features in northeast Spain, are renowned for their rare and threatened flora and fauna, yet their soil C cycle characteristics and soil microbial communities provide additional reasons to monitor the impacts of climate change and protect these vulnerable environments from further anthropogenic disturbances.

Cyanobacteria, as key biocrust components, provide a variety of ecosystem functions in drylands. In this study, to identify whether a cyanobacterial community shift is involved in biocrust succession and whether this is linked to altered ecological functions, we investigated cyanobacterial composition, total carbon and nitrogen contents of biocrusts in the Gurbantunggut Desert. Our findings showed that the biocrust cyanobacteria in the Gurbantunggut desert were mostly filamentous, coexisting with abundant unicellular colonial Chroococcidiopsis. Heterocystous Nostoc, Scytonema and Tolypothrix always represented the majority of biocrust nitrogen-fixing organisms, comprising an average of 92% of the nifH gene reads. Community analysis showed a clear shift in prokaryotic community composition associated with biocrust succession from cyanobacteria- to lichen- and moss-dominated biocrusts, and filamentous non-nitrogen-fixing cyanobacteria-dominated communities were gradually replaced by nitrogen-fixing and unicellular colonial communities. Along the succession, there were concomitant reductions in cyanobacterial relative abundance, whereas Chl-a, total carbon and nitrogen contents increased. Concurrently, distinct carbon and nitrogen stores shifts occurred, implying that the main ecological contribution of cyanobacteria in biocrusts changes from carbon- to nitrogen-fixation along with the succession. Our results suggest that any activity that reverses biocrust succession will influence cyanobacterial community composition and eventually lead to large reductions in soil carbon and nitrogen stores.

Seasonally-flooded várzea forests of Western Amazonia are one of the most productive and biodiverse wetland forests in the world. However, data on their soil CO2 emissions, soil organic matter decomposition rates, and soil C stocks are scarce. This is a concern because hydrological changes are predicted to lead to increases in the height, extent, and duration of seasonal floods, which are likely to have a significant effect on soil C stocks and fluxes. However, with no empirical data, the impact of altered flood regimes on várzea soil C cycles remains uncertain. This study quantified the effects of maximum annual flood height and soil moisture on soil CO2 efflux rate (Rs) and soil organic matter decomposition rate (k) in the várzea forests of Pacaya Samiria National Reserve, Peru. The study was conducted between May and August 2017. The results showed that Rs (10.6–182.7 mg C m−2 h–1) and k (0.016–0.078) varied between and within sites, and were considerably lower than the values reported for other tropical forests. In addition, Rs was negatively affected by flood height (P < 0.01) and soil moisture (P < 0.001), and it decreased with decreasing river levels post flooding (P < 0.001). In contrast, k was not affected by any of the above-mentioned factors. Soil moisture was the dominant factor influencing Rs, and it was significantly affected by maximum flood height, even after the floods had subsided (P < 0.001). Consequently, we concluded that larger floods will likely lead to reduced Rs, whilst k could remain unchanged but with decomposition processes becoming more anaerobic.

Current analyses and predictions of spatially explicit patterns and processes in ecology most often rely on climate data interpolated from standardized weather stations. This interpolated climate data represents long-term average thermal conditions at coarse spatial resolutions only. Hence, many climate-forcing factors that operate at fine spatiotemporal resolutions are overlooked. This is particularly important in relation to effects of observation height (e.g. vegetation, snow and soil characteristics) and in habitats varying in their exposure to radiation, moisture and wind (e.g. topography, radiative forcing or cold-air pooling). Since organisms living close to the ground relate more strongly to these microclimatic conditions than to free-air temperatures, microclimatic ground and near-surface data are needed to provide realistic forecasts of the fate of such organisms under anthropogenic climate change, as well as of the functioning of the ecosystems they live in. To fill this critical gap, we highlight a call for temperature time series submissions to SoilTemp, a geospatial database initiative compiling soil and near-surface temperature data from all over the world. Currently, this database contains time series from 7,538 temperature sensors from 51 countries across all key biomes. The database will pave the way toward an improved global understanding of microclimate and bridge the gap between the available climate data and the climate at fine spatiotemporal resolutions relevant to most organisms and ecosystem processes.

Shrub encroachment is occurring in many of the world's drylands, but its impacts on ecosystem structure and function are still poorly understood. In particular, it remains unclear how shrub encroachment affects dryland soil surfaces, including biological soil crust (biocrust) communities. In this study, soil surfaces (0-1 cm depth) were sampled from areas of Grewia flava shrubs and Eragrostis lehmanniana and Schmidtia kalahariensis grasses in the southwest Kalahari during two different seasons (March and November). Our hypothesis is that the presence of different vegetation cover types (shrubs versus grasses) alters the microbial composition of soil surfaces owing to their contrasting microenvironments. The results showed that more significant differences in microclimate (light, soil surface temperatures) and soil surface microbial communities were observed between shrubs and grasses than between sampling seasons. Based on high-throughput 16S rRNA gene sequencing, our findings showed that approximately one third (33.5%) of the operational taxonomic units (OTUs) occurred exclusively in soil surfaces beneath shrubs. Soil surfaces with biocrusts in grass areas were dominated by the cyanobacteria Microcoleus steenstrupii, whereas the soil surfaces beneath shrubs were dominated by the proteobacteria Microvirga flocculans. Soil surfaces beneath shrubs are associated with reduced cyanobacterial abundance but have higher total carbon and total nitrogen contents compared to biocrusts in grass areas. These findings infer changes in the relative contributions from different sources of carbon and nitrogen (e.g. cyanobacterial and non-cyanobacterial fixation, plant litter, animal activity). The distinctive microbial composition and higher carbon and nitrogen contents in soil surfaces beneath shrubs may provide a positive feedback mechanism promoting shrub encroachment, which helps to explain why the phenomenon is commonly observed to be irreversible. (C) 2020 Elsevier B.V. All rights reserved.

In dryland regions, biological soil crusts (BSCs) have numerous important ecosystem functions. Crust species and functions are, however, highly spatially heterogeneous and remain poorly understood at a range of scales. In this study, chlorophyll fluorescence imaging was used to quantify millimeter-scale patterns in the distribution and activity of photosynthetic organisms in BSCs of different successional stages (including cyanobacterial, lichen, moss three main successional stages and three intermixed transitional stages) from the Tengger Desert, China. Chlorophyll fluorescence images derived from the Imaging PAM (Pulse Amplitude Modulation) showed that with the succession from cyanobacterial to lichen and to moss crusts, crust photosynthetic efficiency (including the maximum and effective photosynthetic efficiency, respectively) and fluorescence coverage increased significantly (P < 0.05), and that increasing photosynthetically active radiation (PAR) reduced the effective photosynthetic efficiency (Yield). The distribution of photosynthetic organisms in crusts determined Fv/Fm (ratio of variable fluorescence to maximum fluorescence) frequency pattern, although the photosynthetic heterogeneity (SHI index) was not significantly different (P > 0.05) between cyanobacterial and moss crusts, and showed a unimodal pattern of Fv/Fm values. In contrast, photosynthetic heterogeneity was significantly higher in lichen, cyanobacteria-moss and lichen-moss crusts (P < 0.05), with a bimodal pattern of Fv/Fm values. Point pattern analysis showed that the distribution pattern of chlorophyll fluorescence varied at different spatial scales and also among the different crust types. These new results provide a detailed (millimeter-scale) insight into crust photosynthetic mechanisms and spatial distribution patterns associated with their community types. Collectively, this information provides an improved theoretical basis for crust maintenance and management in dryland regions.

Drylands contain 25% of the world’s soil organic carbon (SOC), which is controlled by many factors, both abiotic and biotic. Thus, understanding how these factors control SOC concentration can help to design more sustainable land-use practices in drylands aiming to foster and preserve SOC storage, something particularly important to fight ongoing global warming. We use two independent, large-scale databases with contrasting geographic coverage (236 sites in global drylands and 185 sites in Patagonia, Argentina) to evaluate the relative importance of abiotic (precipitation, temperature and soil texture) and biotic (primary productivity) factors as drivers of SOC concentration in drylands at global and regional scales. We found that biotic and abiotic factors had similar effects on SOC concentration across regional and global scales: Maximum temperature and sand content had negative effects, while precipitation and plant productivity exerted positive effects. Our findings provide empirical evidence that increases in temperature and reductions in rainfall, as forecasted by climatic models in many drylands worldwide, promote declines in SOC both directly and indirectly via the reduction in plant productivity. This has important implications for the conservation of drylands under climate change; land management should seek to enhance plant productivity as a tool to offset the negative impact of climate change on SOC storage and on associated ecosystem services.