Jayne Belnap’s research while affiliated with French Geological Survey and other places

The effects of severe drought on the stability of dryland ecosystems are still uncertain and it is unknown whether diversity can buffer changes in systems that are adapted to water-limitation. We investigated the effects of long-term induced drought on the composition and maturity of biological soil crusts (biocrusts), as well as tested the hypothesis that diversity promotes stability using compositional resistance as a measure for ecosystem stability. We surveyed an array of 25 sites in the central Colorado Plateau, USA, that included plots that received ambient precipitation and plots that had experienced eight years of ~35% precipitation reduction. We found that biocrusts can maintain broad compositional integrity after long-term climate disturbance. However, biocrust successional reversal still occurred, with a reduction of later successional constituents and an increase of early successional cyanobacterial cover. Our findings indicate that long-term drought could have major impacts on biocrust community stability.

Understanding the resilience of ecosystems globally is hampered by the complex and interacting drivers of change characteristic of the Anthropocene. This is true for drylands of the western US, where widespread alteration of disturbance regimes and spread of invasive non-native species occurred with westward expansion during the 1800s, including the introduction of domestic livestock and spread of Bromus tectorum, an invasive non-native annual grass. In addition, this region has experienced a multi-decadal drought not seen for at least 1200 years with potentially large and interacting impacts on native plant communities. Here, we present 24 years of twice-annual plant cover monitoring (1997-2021) from a semiarid grassland never grazed by domestic livestock but subject to a patchy invasion of B. tectorum beginning in ~1994, compare our findings to surveys done in 1967, and examine potential climate drivers of plant community changes. We found a significant warming trend in the study area, with more than 75% of study year temperatures being warmer than average (1966-2021). We observed a native perennial grass community with high resilience to climate forcings with cover values like those in 1967. In invaded patches, B. tectorum cover was greatest in the early years of this study (1997-2001; ~20%-40%) but was subsequently constrained by climate and subtle variation in soils, with limited evidence of long-term impacts to native vegetation, contradicting earlier studies. Our ability to predict year-to-year variation in functional group and species cover with climate metrics varied, with a 12-month integrated index and fall and winter patterns appearing most important. However, declines to near zero live cover in recent years in response to regional drought intensification leave questions regarding the resiliency of intact grasslands to ongoing aridification and whether the vegetation observations reported here may be a leading indicator of impending change in this protected ecosystem.

Global dryland vegetation communities will likely change as ongoing drought conditions shift regional climates towards a more arid future. Additional aridification of drylands can impact plant and ground cover, biogeochemical cycles, and plant-soil feedbacks, yet how and when these crucial ecosystem components will respond to drought intensification requires further investigation. Using a long-term precipitation reduction experiment (35% reduction) conducted across the Colorado Plateau and spanning ten years into a 20+ year regional megadrought, we explored how vegetation cover, soil conditions, and growing season nitrogen (N) availability are impacted by drying climate conditions. We observed large declines for all dominant plant functional types (C3 and C4 grasses and C3 and C4 shrubs) across measurement period, both in the drought treatment and control plots, likely due to ongoing regional megadrought conditions. In experimental drought plots, we observed less plant cover, less biological soil crust cover, warmer and drier soil conditions, and more soil resin-extractable N compared to the control plots. Observed increases in soil N availability were best explained by a negative correlation with plant cover regardless of treatment, suggesting that declines in vegetation N uptake may be driving increases in available soil N. However, in ecosystems experiencing long-term aridification, increased N availability may ultimately result in N losses if soil moisture is consistently too dry to support plant and microbial N immobilization and ecosystem recovery. These results show dramatic, worrisome declines in plant cover with long-term drought. Additionally, this study highlights that more plant cover losses are possible with further drought intensification, and underscore that, in addition to large drought effects on aboveground communities, drying trends drive significant changes to critical soil resources such as N availability, all of which could have long-term ecosystem impacts for drylands.

Responses of terrestrial ecosystems to climate change have been explored in many regions worldwide. While continued drying and warming may alter process rates and deteriorate the state and performance of ecosystems, it could also lead to more fundamental changes in the mechanisms governing ecosystem functioning. Here we argue that climate change will induce unprecedented shifts in these mechanisms in historically wetter climatic zones, towards mechanisms currently prevalent in dry regions, which we refer to as ‘dryland mechanisms’. We discuss 12 dryland mechanisms affecting multiple processes of ecosystem functioning, including vegetation development, water flow, energy budget, carbon and nutrient cycling, plant production and organic matter decomposition. We then examine mostly rare examples of the operation of these mechanisms in non-dryland regions where they have been considered irrelevant at present. Current and future climate trends could force microclimatic conditions across thresholds and lead to the emergence of dryland mechanisms and their increasing control over ecosystem functioning in many biomes on Earth. In drylands, there are unique mechanisms that influence multiple ecosystem processes. In this Perspective, the authors identify these dryland mechanisms and show that they could become more important in non-dryland regions or areas that will become drier in the future. *** FOR ONLINE VIEWING: https://rdcu.be/cSmAD

Biological soil crusts (biocrusts) cover ~12% of the global land surface. They are formed by an intimate association between soil particles, photoautotrophic and heterotrophic organisms, and they effectively stabilize the soil surface of drylands. Quantitative information on the impact of biocrusts on the global cycling and climate effects of aeolian dust, however, is not available. Here, we combine the currently limited experimental data with a global climate model to investigate the effects of biocrusts on regional and global dust cycling under current and future conditions. We estimate that biocrusts reduce the global atmospheric dust emissions by ~60%, preventing the release of ~0.7 Pg dust per year. Until 2070, biocrust coverage is expected to be severely reduced by climate change and land-use intensification. The biocrust loss will cause an increased dust burden, leading to a reduction of the global radiation budget of around 0.12 to 0.22 W m⁻², corresponding to about 50% of the total direct forcing of anthropogenic aerosols. This biocrust control on dust cycling and its climate impacts have important implications for human health, biogeochemical cycling and the functioning of the ecosystems, and thus should be considered in the modelling, mitigation and management of global change.

Studies of biological soil crusts (biocrusts) have proliferated over the last few decades. The biocrust literature has broadened, with more studies assessing and describing the function of a variety of biocrust communities in a broad range of biomes and habitats and across a large spectrum of disciplines, and also by the incorporation of biocrusts into global perspectives and biogeochemical models. As the number of biocrust researchers increases, along with the scope of soil communities defined as ‘biocrust’, it is worth asking whether we all share a clear, universal, and fully articulated definition of what constitutes a biocrust. In this review, we synthesize the literature with the views of new and experienced biocrust researchers, to provide a refined and fully elaborated definition of biocrusts. In doing so, we illustrate the ecological relevance and ecosystem services provided by them. We demonstrate that biocrusts are defined by four distinct elements: physical structure, functional characteristics, habitat, and taxonomic composition. We describe outgroups, which have some, but not all, of the characteristics necessary to be fully consistent with our definition and thus would not be considered biocrusts. We also summarize the wide variety of different types of communities that fall under our definition of biocrusts, in the process of highlighting their global distribution. Finally, we suggest the universal use of the Belnap, Büdel & Lange definition, with minor modifications: Biological soil crusts (biocrusts) result from an intimate association between soil particles and differing proportions of photoautotrophic (e.g. cyanobacteria, algae, lichens, bryophytes) and heterotrophic (e.g. bacteria, fungi, archaea) organisms, which live within, or immediately on top of, the uppermost millimetres of soil. Soil particles are aggregated through the presence and activity of these often extremotolerant biota that desiccate regularly, and the resultant living crust covers the surface of the ground as a coherent layer. With this detailed definition of biocrusts, illustrating their ecological functions and widespread distribution, we hope to stimulate interest in biocrust research and inform various stakeholders (e.g. land managers, land users) on their overall importance to ecosystem and Earth system functioning.

Significance Across many global drylands, biocrusts form a protective barrier on the soil surface and fill many critical roles in these harsh yet fragile environments. Previous short-term research suggests that climate change and invasive plant introduction can damage and alter biocrust communities, yet few long-term observations exist. Using a globally unique long-term record of continuous biocrust surveys from a rare never-grazed, protected grassland on the US Colorado Plateau, we found lichen species diversity and cover to be negatively correlated with increasing summer air temperatures, while moss species showed more sensitivity to variation in precipitation and invasive grass cover. These results suggest that dryland systems may be at a critical tipping point where ongoing warming could result in biological soil crust degradation.

Global concerns for desertification have focused on the slow recovery of extensive and expanding drylands following disturbance, which may be exacerbated by climate change. Biological soil crusts (biocrusts) are photosynthetic soil communities found in drylands worldwide, which are central to the stability and resilience of dryland ecosystems, but vulnerable to global change. Here we use multiple decade-long experiments to investigate the consequences of climate and land-use change on biocrusts and soil stability. Biocrusts recovered rapidly under ambient temperatures but warming interacted with the precipitation disturbance to halt recovery. Moreover, warming alone caused losses of mosses, lichens and soil stability. Our results present a new mechanism contributing to land degradation in drylands whereby warming drives a state shift in biocrust communities, which degrades soil stability. The synergistic effects of climate and land-use change co-occur globally and our results support projections of increased desertification and lowered dryland resilience under warming.

Biological soil crusts (biocrusts) cover the soil surface of global drylands and interact with vascular plants. Biocrusts may influence the availability and nature of safe sites for plant recruitment and the susceptibility of an area to invasion by non‐native species. Therefore, to investigate the potential role of biocrusts in invasive species management, we sought to determine whether native and non‐native grass recruitments in two North American deserts were differentially affected by biocrusts. We conducted a series of coordinated experiments in field, semi‐controlled, and controlled environment settings in the Colorado Plateau and Sonoran Desert using contrasting biocrust and grass functional types. Experiments in field environments focused on early establishment of grass seedlings whereas controlled environment experiments focused on seedling emergence. Within each experiment, we compared responses (frequency, magnitude, and timing of emergence/establishment) both across species (biocrust types pooled) and across species and levels of biocrust development. Native grasses varied by experiment and included Aristida purpurea, A. purpurea var. longiseta, Bouteloua gracilis, and Vulpia octoflora. Emergence of non‐native Bromus tectorum was similar to that of native grasses on the Colorado Plateau. Differences in emergence of native vs. non‐native grasses in the Sonoran Desert were species‐ and response‐specific. Emergence of the non‐native Bromus rubens was comparable to that of native grasses whereas emergence frequency and magnitude of the non‐native Pennisetum ciliare was lower compared with two of four native species. Within a grass species, emergence was higher and faster on bare soil compared with biocrusts in the Sonoran Desert semi‐controlled and greenhouse environment experiments. However, the pattern was not consistent across other experiments. When comparing across Colorado Plateau and Sonoran Desert biocrusts in greenhouse experiments, we found that emergence of native grasses was higher on Colorado Plateau biocrusts. Based on the lack of consistent results across our experiments, grass recruitment on biocrusts appears to be driven more by species‐specific traits than species provenance. Our greenhouse experiments suggest that biocrust topographic relief is an important safe site trait influencing plant recruitment.

Dryland ecosystems are sensitive to perturbations and generally slow to recover post disturbance. The microorganisms residing in dryland soils are especially important as they contribute to soil structure and nutrient cycling. Disturbance can have particularly strong effects on dryland soil structure and function, yet the natural resistance and recovery of the microbial components of dryland soils has not been well documented. In this study, the recovery of surface soil bacterial communities from multiple physical and environmental disturbances is assessed. Samples were collected from three field sites in the vicinity of Moab, UT, United States, 6 to 7 years after physical and climate disturbance manipulations had been terminated, allowing for the assessment of community recovery. Additionally, samples were collected in a transect that included three habitat patches: the canopy zone soils under the dominant shrubs, the interspace soils that are colonized by biological soil crusts, and edge soils at the plot borders. Field site and habitat patch were significant factors structuring the bacterial communities, illustrating that sites and habitats harbored unique soil microbiomes. Across the different sites and disturbance treatments, there was evidence of significant bacterial community recovery, as bacterial biomass and diversity were not significantly different than control plots. There was, however, a small number of 16S rRNA gene amplicon sequence variants that distinguished particular treatments, suggesting that legacy effects of the disturbances still remained. Taken together, these data suggest that dryland bacterial communities may possess a previously unappreciated potential to recover within years of the original disturbance.