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Land use practices and climate change have driven substantial soil degradation across global drylands, impacting ecosystem functions and human livelihoods. Biological soil crusts, a common feature of dryland ecosystems, are under extensive exploration for their potential to restore the stability and fertility of degraded soils through the development of inoculants. However, stressful abiotic conditions often result in the failure of inoculation-based restoration in the field and may hinder the long-term success of biocrust restoration efforts. Taking an assisted migration approach, we cultivated biocrust inocula sourced from multiple hot-adapted sites (Mojave and Sonoran Deserts) in an outdoor facility at a cool desert site (Colorado Plateau). In addition to cultivating inoculum from each site, we created an inoculum mixture of biocrust from the Mojave Desert, Sonoran Desert, and Colorado Plateau. We then applied two habitat amelioration treatments to the cultivation site (growth substrate and shading) to enhance soil stability and water availability and reduce UV stress. Using marker gene sequencing, we found that the cultivated mixed inoculum comprised both local- and hot-adapted cyanobacteria at the end of cultivation but had similar cyanobacterial richness as each unmixed inoculum. All cultivated inocula had more cyanobacterial 16S rRNA gene copies and higher cyanobacterial richness when cultivated with a growth substrate and shade. Our work shows that it is possible to field cultivate biocrust inocula sourced from different deserts, but that community composition shifts toward that of the cultivation site unless habitat amelioration is employed. Future assessments of the function of a mixed inoculum in restoration and its resilience in the face of abiotic stressors are needed to determine the relative benefit of assisted migration compared to the challenges and risks of this approach.

Up to 35% of global drylands have experienced degradation due to anthropogenic impacts, including physical disturbances like trampling and soil removal. These physical disturbances can result in the loss of soil communities known as biological soil crusts (biocrusts) and the important functions they provide, such as soil stability and fertility. The reestablishment of biocrust organisms after disturbance is determined by many factors, including propagule availability, climate, and vascular plant community structure. The role of these factors in natural recovery may be intensified by the extent (or size) of a disturbance. For example, large disturbances can result in reduced propagule availability or enhanced erosion, which impact both the dispersal and establishment of biocrust organisms on disturbed soils, leading to a slower natural recovery. To test how disturbance extent impacts biocrust's natural recovery, we installed four disturbance extents by completely removing biocrust from the mineral soil in plots ranging from 0.01 m 2 to 1 m 2 and measured productivity and erosion resistance. We found that small disturbance extents did not differ in chlorophyll a content, total exopolysaccharide content, or soil stability after 1.5 years of natural recovery. However, the concentration of glycocalyx exopolysaccharide was higher in the smallest disturbances after the recovery period. Our results indicate that disturbances <1 m 2 in scale recover at similar rates, with soil stability returning to high levels in just a few years after severe disturbance. Our findings align with prior work on biocrust natural recovery in drylands and highlight the opportunity for future work to address (1) cyanobacteria, moss, and lichen propagule dispersal; (2) rates and mechanisms of biocrust succession; and (3) the role of wind or water in determining biocrust colonization patterns as compared to lateral growth., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2023 Jech, Havrilla and Barger.)

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., (© 2022 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Biological soil crusts (biocrusts) are a complex community of algae, cyanobacteria, lichens, bryophytes, and assorted bacteria, fungi, archaea, and bacteriophages that colonize the soil surface. Biocrusts are particularly common in drylands and are found in arid and semiarid ecosystems worldwide. While diminutive in size, biocrusts often cover large terrestrial areas, provide numerous ecosystem benefits, enhance biodiversity, and are found in multiple configurations and assemblages across different climate and disturbance regimes. Biocrusts have been a focus of many ecologists, especially those working in semiarid and arid lands, as biocrusts are foundational community members, play fundamental roles in ecosystem processes, and offer rare opportunities to study biological interactions at small and large spatial scales. Due to these same characteristics, biocrusts have the potential to serve as an excellent teaching tool. The purpose of this paper is to demonstrate the utility of biocrust communities as a model system in science education. Functioning as portable, dynamic mini ecosystems, biocrusts can be used to teach about organisms, biodiversity, biotic interactions, abiotic controls, ecosystem processes, and even global change, and can be easy to use in nearly every classroom setup. For example, education principles, such as evolution and adaptation to stress, or structure and function (patterns and processes) can be applied by bringing biocrusts into the classroom as a teaching tool. In addition, discussing the utility of biocrusts in the classroom - including theory, hypothesis testing, experimentation, and hands-on learning - this document also provides tips and resources for developing education tools and activities geared toward impactful learning., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Faist, Antoninka, Barger, Bowker, Chaudhary, Havrilla, Huber-Sannwald, Reed and Weber.)

Fairy circles are striking regularly sized and spaced, bare circles surrounded by Stipagrostis grasses that occur over thousands of square kilometres in Namibia. The mechanisms explaining their origin, shape, persistence and regularity remain controversial. One hypothesis for the formation of vegetation rings is based on the centrifugal expansion of a single individual grass plant, via clonal growth and die-back in the centre. Clonality could explain FC origin, shape and long-term persistence as well as their regularity, if one clone competes with adjacent clones. Here, we show that for virtually all tested fairy circles the periphery is not exclusively made up of genetically identical grasses, but these peripheral grasses belong to more than one unrelated genet. These results do not support a clonal explanation for fairy circles. Lack of clonality implies that a biological reason for their origin, shape and regularity must emerge from competition between near neighbor individuals within each fairy circle. Such lack of clonality also suggests a mismatch between longevity of fairy circles versus their constituent plants. Furthermore, our findings of lack of clonality have implications for some models of spatial patterning of fairy circles that are based on self-organization.

Biological soil crusts (biocrusts) are topsoil communities formed by cyanobacteria or other microbial primary producers and are typical of arid and semiarid environments. Biocrusts promote a range of ecosystem services, such as erosion resistance and soil fertility, but their degradation by often anthropogenic disturbance brings about the loss of these services. This has prompted interest in developing restoration techniques. One approach is to source biocrust remnants from the area of interest for scale-up cultivation in a microbial "nursery" that produces large quantities of high-quality inoculum for field deployment. However, growth dynamics and the ability to reuse the produced inoculum for continued production have not been assessed. To optimize production, we followed nursery growth dynamics of biocrusts from cold (Great Basin) and hot (Chihuahuan) deserts. Peak phototrophic biomass was attained between 3 and 7 weeks in cold desert biocrusts and at 12 weeks in those from hot deserts. We also reused the resultant biocrust inoculum to seed successive incubations, tracking both phototroph biomass and cyanobacterial community structure using 16S rRNA gene amplicon sequencing. Hot desert biocrusts showed little to no viability upon reinoculation, while cold desert biocrusts continued to grow, but at the expense of progressive shifts in species composition. This leads us to discourage the reuse of nursery-grown inoculum. Surprisingly, growth was highly variable among replicates, and overall yields were low, a fact that we attribute to the demonstrable presence of virulent and stochastically distributed but hitherto unknown cyanobacterial pathogens. We provide recommendations to avoid pathogen incidence in the process. IMPORTANCE Biocrust communities provide important ecosystem services for arid land soils, such as soil surface stabilization promoting erosion resistance and contributing to overall soil fertility. Anthropogenic degradation to biocrust communities (through livestock grazing, agriculture, urban sprawl, and trampling) is common and significant, resulting in a loss of those ecosystem services. Losses impact both the health of the native ecosystem and the public health of local populations due to enhanced dust emissions. Because of this, approaches for biocrust restoration are being developed worldwide. Here, we present optimization of a nursery-based approach to scaling up the production of biocrust inoculum for field restoration with respect to temporal dynamics and reuse of biological materials. Unexpectedly, we also report on complex population dynamics, significant spatial variability, and lower than expected yields that we ascribe to the demonstrable presence of cyanobacterial pathogens, the spread of which may be enhanced by some of the nursery production standard practices., (Copyright © 2019 Bethany et al.)

Biological soil crusts (biocrusts) are slow-growing, phototroph-based microbial assemblages that develop on the topsoils of drylands. Biocrusts help maintain soil fertility and reduce erosion. Because their loss through human activities has negative ecological and environmental health consequences, biocrust restoration is of interest. Active soil inoculation with biocrust microorganisms can be an important tool in this endeavor. We present a culture-independent, two-step process to grow multispecies biocrusts in open greenhouse nursery facilities, based on the inoculation of local soils with local biocrust remnants and incubation under seminatural conditions that maintain the essence of the habitat but lessen its harshness. In each of four U.S. Southwest sites, we tested and deployed combinations of factors that maximized growth (gauged as chlorophyll a content) while minimizing microbial community shifts (assessed by 16S rRNA sequencing and bioinformatics), particularly for crust-forming cyanobacteria. Generally, doubling the frequency of natural wetting events, a 60% reduction in sunlight, and inoculation by slurry were optimal. Nutrient addition effects were site specific. In 4 months, our approach yielded crusts of high inoculum quality reared on local soil exposed to locally matched climates, acclimated to desiccation, and containing communities minimally shifted in composition from local ones. Our inoculum contained abundant crust-forming cyanobacteria and no significant numbers of allochthonous phototrophs, and it was sufficient to treat ca. 6,000 m 2 of degraded dryland soils at 1 to 5% of the typical crust biomass concentration, having started from a natural crust remnant as small as 6 to 30 cm 2 IMPORTANCE: Soil surface crusts can protect dryland soils from erosion, but they are often negatively impacted by human activities. Their degradation causes a loss of fertility, increased production of fugitive dust and intensity of dust storms with associated traffic problems, and provokes general public health hazards. Our results constitute an advance in the quest to actively restore biological soil covers by providing a means to obtain high-quality inoculum within a reasonable time (a few months), thereby allowing land managers to recover essential, but damaged, ecosystem services in a sustainable, self-perpetuating way as provided by biocrust communities., (Copyright © 2017 American Society for Microbiology.)

Recent droughts and increasing temperatures have resulted in extensive tree mortality across the globe. Understanding the environmental controls on tree regeneration following these drought events will allow for better predictions of how these ecosystems may shift under a warmer, drier climate. Within the widely distributed piñon-juniper woodlands of the southwestern USA, a multiyear drought in 2002-2004 resulted in extensive adult piñon mortality and shifted adult woodland composition to a juniper-dominated, more savannah-type ecosystem. Here, we used pre- (1998-2001) and 10-year post- (2014) drought stand structure data of individually mapped trees at 42 sites to assess the effects of this drought on tree regeneration across a gradient of environmental stress. We found declines in piñon juvenile densities since the multiyear drought due to limited new recruitment and high (>50%) juvenile mortality. This is in contrast to juniper juvenile densities, which increased over this time period. Across the landscape, piñon recruitment was positively associated with live adult piñon densities and soil available water capacity, likely due to their respective effects on seed and water availability. Juvenile piñon survival was strongly facilitated by certain types of nurse trees and shrubs. These nurse plants also moderated the effects of environmental stress on piñon survival: Survival of interspace piñon juveniles was positively associated with soil available water capacity, whereas survival of nursed piñon juveniles was negatively associated with perennial grass cover. Thus, nurse plants had a greater facilitative effect on survival at sites with higher soil available water capacity and perennial grass cover. Notably, mean annual climatic water deficit and elevation were not associated with piñon recruitment or survival across the landscape. Our findings reveal a clear shift in successional trajectories toward a more juniper-dominated woodland and highlight the importance of incorporating biotic interactions and soil properties into species distribution modeling approaches., (© 2015 John Wiley & Sons Ltd.)

National fuel-reduction programs aim to reduce the risk of wildland fires to human communities and to restore forest and rangeland ecosystems to resemble their historical structure, function, and diversity. There are a number of factors, such as seed bank dynamics, post-treatment climate, and herbivory, which determine whether this latter goal may be achieved. Here, we examine the short-term (2 years) vegetation response to fuel-reduction treatments (mechanical mastication, broadcast burn, and pile burn) and seeding of native grasses on understory vegetation in an upland piñon-juniper woodland in southeast Utah. We also examine how wildlife herbivory affects the success of fuel-reduction treatments. Herbaceous cover increased in response to fuel-reduction treatments in all seeded treatments, with the broadcast burn and mastication having greater increases (234 and 160 %, respectively) in herbaceous cover than the pile burn (32 %). In the absence of seeding, herbaceous cover only increased in the broadcast burn (32 %). Notably, fuel-reduction treatments, but not seeding, strongly affected herbaceous plant composition. All fuel-reduction treatments increased the relative density of invasive species, especially in the broadcast burn, which shifted the plant community composition from one dominated by perennial graminoids to one dominated by annual forbs. Herbivory by wildlife reduced understory plant cover by over 40 % and altered plant community composition. If the primary management goal is to enhance understory cover while promoting native species abundance, our study suggests that mastication may be the most effective treatment strategy in these upland piñon-juniper woodlands. Seed applications and wildlife exclosures further enhanced herbaceous cover following fuel-reduction treatments.

Widespread piñon (Pinus edulis) mortality occurred across the southwestern USA during 2002-2003 in response to drought and bark beetle infestations. Given the recent mortality and changes in regional climate over the past several decades, there is a keen interest in post-mortality regeneration dynamics in piñon-juniper woodlands. Here, we examined piñon and Utah juniper (Juniperus osteosperma) recruitment at 30 sites across southwestern Colorado, USA that spanned a gradient of adult piñon mortality levels (10-100%) to understand current regeneration dynamics. Piñon and juniper recruitment was greater at sites with more tree and shrub cover. Piñon recruitment was more strongly facilitated than juniper recruitment by trees and shrubs. New (post-mortality) piñon recruitment was negatively affected by recent mortality. However, mortality had no effect on piñon advanced regeneration (juveniles established pre-mortality) and did not shift juvenile piñon dominance. Our results highlight the importance of shrubs and juniper trees for the facilitation of piñon establishment and survival. Regardless of adult piñon mortality levels, areas with low tree and shrub cover may become increasingly juniper dominated as a result of the few suitable microsites for piñon establishment and survival. In areas with high piñon mortality and high tree and shrub cover, our results suggest that piñon is regenerating via advanced regeneration., (© 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.)

Background: Increases in the spatial extent and density of woody plants relative to herbaceous species have been observed across many ecosystems. These changes can have large effects on ecosystem carbon stocks and therefore are of interest for regional and national carbon inventories and for potential carbon sequestration or management activities. However, it is challenging to estimate the effect of woody plant encroachment on carbon because aboveground carbon stocks are very heterogeneous spatially and belowground carbon stocks exhibit complex and variable responses to changing plant cover. As a result, estimates of carbon stock changes with woody plant cover remain highly uncertain. In this study, we use a combination of plot- and remote sensing-based techniques to estimate the carbon impacts of piñon and juniper (PJ) encroachment in SE Utah across a variety of spatial scales with a specific focus on the role of spatial heterogeneity in carbon estimates., Results: At a plot scale (300 m2) areas piñon juniper (PJ) encroached areas had 0.26 kg C m-2 less understory vegetation carbon compared to un-encroached sites. This lower amount of carbon was offset by an average of 1.82 kg C m-2 higher carbon in PJ vegetation and 0.50 kg m-2 of C in PJ surface-litter carbon. Soil mineral carbon stocks were unaffected by woody plant cover and density. Aboveground carbon stocks were highly dependent on PJ vegetation density. At a 300 m2 plot-scale, plots with low and high density of PJ forest had 1.40 kg C m-2 and 3.69 kg m-2 more carbon than the un-encroached plot. To examine how these 300 m2 variations influence landscape scale C estimates, historical and contemporary aerial photos were analyzed to develop forest density maps in order to estimate above ground PJ associated C stock changes in a 25 ha area. This technique yielded an average estimate of 1.43 kg m-2 of C accumulation with PJ encroachment. Combining this estimate with analysis of tree growth increments from dendrochronologies, we estimate that these PJ stands are accumulating aboveground C at an annual rate of 0.02 kg C m-2 with no slowing of this rate in healthy PJ. This result is in contrast to what has been observed in large areas of drought related PJ mortality, where C accumulation has ceased., Conclusions: These results illustrate that the encroachment of PJ forests in SE Utah over the last century has resulted in a large (and ongoing) accumulation of carbon in PJ trees and surface litter. However, the magnitude of the increase depends to on the density of vegetation across the landscape and the health of forest stands. Both management activities that remove forest carbon and forest mortality due to drought or wildfire have the potential to quickly reverse the multi-decadal accumulation of carbon in these stands.

Causes of over-dispersed barren "fairy circles" that are often surrounded by ca. 0.5 m tall peripheral grasses in a matrix of shorter (ca. 0.2 m tall) grasses in Namibian grasslands remain mysterious. It was hypothesized that the fairy circles are the consequence of self-organizing spatial vegetation patterning arising from resource competition and facilitation. We examined the edaphic properties of fairy circles and variation in fairy circle size, density and landscape occupancy (% land surface) with edaphic properties and water availability at a local scale (<50 km) and with climate and vegetation characteristics at a regional scale. Soil moisture in the barren fairy circles declines from the center towards the periphery and is inversely correlated with soil organic carbon, possibly indicating that the peripheral grass roots access soil moisture that persists into the dry season within fairy circles. Fairy circle landscape occupancy is negatively correlated with precipitation and soil [N], consistent with fairy circles being the product of resource-competition. Regional fairy circle presence/absence is highly predictable using an empirical model that includes narrow ranges of vegetation biomass, precipitation and temperature seasonality as predictor variables, indicating that fairy circles are likely a climate-dependent emergent phenomenon. This dependence of fairy circle occurrence on climate explains why fairy circles in some locations may appear and disappear over time. Fairy circles are only over-dispersed at high landscape occupancies, indicating that inter-circle competition may determine their spacing. We conclude that fairy circles are likely to be an emergent arid-grassland phenomenon that forms as a consequence of peripheral grass resource-competition and that the consequent barren circle may provide a resource-reservoir essential for the survival of the larger peripheral grasses and provides a habitat for fossicking fauna.

Over the past several decades, the expansion and thickening of woodlands in the western United States has caused a range of ecological changes. Woody expansion often leads to increases in soil organic matter (SOM) pools with implications for both biogeochemical cycling and ecological responses to management strategies aimed at restoration of rangeland ecosystems. Here we directly measure C and N stocks and use simple non-steady-state models to quantify the dynamics of soil C accumulation under and around trees of varied ages in southern Utah woodlands. In the two pinyon-juniper forests of Grand Staircase Escalante National Monument studied here, we found approximately 3 kg C/m2 and approximately 0.12 kg N/m2 larger C and N stocks in soils under pinyon canopies compared to interspace sites. These apparent increases in soil C and N stocks under woody plant species were dominated by elevated SOM in the surface 10 cm of soil, particularly within non-mineral-associated organic fractions. The most significant accumulation of C was in the >850 microm fraction, which had an estimated C residence time of <20 yr. Rates of carbon accumulation following pinyon-juniper expansion appear to be dominated by changes in this fast-cycling surface soil fraction. In contrast, we found that after separating >850 microm organic matter from the remaining light fraction (LF), C had residence times of approximately 400 yr and mineral-associated (MA) soil C had residence times of approximately 600 yr. As a result, we calculate that input rates to the LF and MA pools to be 10 +/- 1 and 0.68 +/- 0.15 g m(-2) yr(-1) (mean +/- SE), respectively. These findings suggest that one consequence of management activities aimed at the reduction of pinyon-juniper biomass may be a relatively rapid loss of soil C and N pools associated with the >850 microm fraction. The temporal dynamics of the <850 microm pools suggest that carbon and nitrogen continue to accumulate in these fractions, albeit at very slow rates, and suggest that multidecadal storage of C following tree recruitment is limited to relatively small, subsurface fractions of the total soil C pool.

Regional, high-resolution mapping of vegetation cover and biomass is central to understanding changes to the terrestrial carbon (C) cycle, especially in the context of C management. The third most extensive vegetation type in the United States is pinyon-juniper (P-J) woodland, yet the spatial patterns of tree cover and aboveground biomass (AGB) of P-J systems are poorly quantified. We developed a synoptic remote-sensing approach to scale up pinyon and juniper projected cover (hereafter "cover") and AGB field observations from plot to regional levels using fractional photosynthetic vegetation (PV) cover derived from airborne imaging spectroscopy and Landsat satellite data. Our results demonstrated strong correlations (P < 0.001) between field cover and airborne PV estimates (r2 = 0.92), and between airborne and satellite PV estimates (r2 = 0.61). Field data also indicated that P-J AGB can be estimated from canopy cover using a unified allometric equation (r2 = 0.69; P < 0.001). Using these multiscale cover-AGB relationships, we developed high-resolution, regional maps of P-J cover and AGB for the western Colorado Plateau. The P-J cover was 27.4% +/- 9.9% (mean +/- SD), and the mean aboveground woody C converted from AGB was 5.2 +/- 2.0 Mg C/ha. Combining our data with the southwest Regional Gap Analysis Program vegetation map, we estimated that total contemporary woody C storage for P-J systems throughout the Colorado Plateau (113 600 km2) is 59.0 +/- 22.7 Tg C. Our results show how multiple remote-sensing observations can be used to map cover and C stocks at high resolution in drylands, and they highlight the role of P-J ecosystems in the North American C budget.