Biocrust Communities Research Articles

The Bacterial Microbiome Associated With Arid Biocrusts and the Biogeochemical Influence of Biocrusts Upon the Underlying Soil.

Biocrusts are aggregated crusts that exist on the soil surface of arid environments. They are complex microbial communities comprised of cyanobacteria, lichens, mosses, algae and fungi. Recently, biocrusts have gained significant attention due to their ubiquitous distribution and likely important ecological roles, including soil stabilization, soil moisture retention, carbon (C) and nitrogen (N) fixation, as well as microbial engineers for semi-arid ecosystem restoration. Here, we collected three co-occurring types of biocrust (Cyanobacterial crust, Crustose lichen, and Foliose lichen) and their underlying soil from arid zones within Western Australia. Bacterial microbiome composition was determined through 16S rRNA gene amplicon sequencing to assess the extent of microbiome selection within the crusts versus underlying soil and biogeochemical measures performed to determine whether the crusts had significant impact upon the underlying soil for nutrient input. We determined that the bacterial communities of native biocrusts are distinct from those in their underlying soil, where dominant bacterial taxa differed according to crust morphologies. δ15N revealed that N-fixation appeared most evident in Foliose lichen crust (1.73 ± 1.04‰). Consequently, depending upon the crust type, biocrusts contained higher concentrations of organic C (2 to 50 times), total N (4 to 16 times) and available ammonium (2 to 4 times), though this enrichment did not extend to the soils underneath them. These findings demonstrate that biocrust communities are seemingly islands of biological activity in an arid landscape, uniquely different from their surrounding and underlying soil.

Biocrust landscape‐scale spatial distribution is strongly controlled by terrain attributes: Topographic thresholds for colonization in a semiarid badland system

AbstractBiological soil crust, or biocrust communities, are the dominating life form in many extreme habitats, such as arid and semiarid badlands, where water scarcity and highly erodible substrates limit vegetation cover. While climate, soil and biotic factors have been described as environmental filters influencing biocrust distribution in such biomes, little is known about the effect of terrain attributes on creating specific microhabitats that promote or restrict biocrust colonization. This study aimed to identify the main terrain attributes controlling biocrust distribution in the driest badland system in Europe, the Tabernas Badlands (SE Spain). To do this, we analysed the influence of different terrain attributes related to landscape stability and microclimate formation on the spatial distribution of lichen and cyanobacteria, using field measurements and topographical information from a LiDAR survey. Our results showed that the spatial distribution of cyanobacteria‐dominated biocrusts, which are physiologically and morphologically adapted to extreme drought and high UVA radiation, was mostly associated with areas of high potential incoming solar radiation. The exception was bare south‐aspect hillslopes with very high sediment transport potential, where bare physically crusted soils were the dominant ground cover. Lichen‐dominated biocrusts, in contrast, colonized near the top of north‐aspect hillslopes, characterized by low potential incoming solar radiation and potential evapotranspiration, and their cover decreased downstream, as conditions became good enough for vascular plants. © 2019 John Wiley & Sons, Ltd. © 2019 John Wiley & Sons, Ltd.

Cold Acclimation Improves the Desiccation Stress Resilience of Polar Strains of Klebsormidium (Streptophyta).

Biological soil crusts (BSCs) are complex communities of autotrophic, heterotrophic, and saprotrophic (micro)organisms. In the polar regions, these biocrust communities have essential ecological functions such as primary production, nitrogen fixation, and ecosystem engineering while coping with extreme environmental conditions (temperature, desiccation, and irradiation). The microalga Klebsormidium is commonly found in BSCs all across the globe. The ecophysiological resilience of various Klebsormidium species to desiccation and other stresses has been studied intensively. Here we present the results of transcriptomic analyses of two different Klebsormidium species, K. dissectum and K. flaccidum, isolated from Antarctic and Arctic BSCs. We performed desiccation stress experiments at two different temperatures mimicking fluctuations associated with global change. Cultures grown on agar plates were desiccated on membrane filters at 10% relative air humidity until the photosynthetic activity as reflected in the effective quantum yield of photosystem II [Y(II)] ceased. For both species, the response to dehydration was much faster at the higher temperature. At the transcriptome level both species responded more strongly to the desiccation stress at the higher temperature suggesting that adaptation to cold conditions enhanced the resilience of both algae to desiccation stress. Interestingly, the two different species responded differently to the applied desiccation stress with respect to the number as well as function of genes showing differential gene expression. The portion of differentially expressed genes shared between both taxa was surprisingly low indicating that both Klebsormidium species adapted independently to the harsh conditions of Antarctica and the Arctic, respectively. Overall, our results indicate that environmental acclimation has a great impact on gene expression and the response to desiccation stress in Klebsormidium.

A practical guide to measuring functional indicators and traits in biocrusts

Biocrusts are multifunctional communities that are increasingly being used to restore degraded or damaged ecosystems. Concurrently, restoration science is shifting away from the use of purely structural metrics, such as relative abundance, to more functional approaches. Although biocrust restoration technology is advancing, there is a lack of readily available information on how to monitor biocrust functioning and set appropriate restoration goals. We therefore compiled a selection of 22 functional indicators that can be used to monitor biocrust functions, such as CO2 exchange as an indicator of productivity or soil aggregate stability as a proxy for erosion resistance. We describe the functional importance of each indicator and the available protocols with which it may be measured. The majority of indicators can be measured as a functional trait of species by using patches of biocrust or cultures that contain only one species. Practitioners wishing to track the multifunctionality of an entire biocrust community would be advised to choose one indicator from each broad functional group (erosion resistance, nutrient accumulation, productivity, energy balance, hydrology), whereas a targeted approach would be more appropriate for projects with a key function of interest. Because predisturbance data are rarely available for biocrust functions, restoration goals can be based on a closely analogous site, literature values, or an expert elicitation process. Finally, we advocate for the establishment of a global trait database for biocrusts, which would reduce the damage resulting from repeated sampling, and provide a wealth of future research opportunities.

Optimizing the Production of Nursery-Based Biological Soil Crusts for Restoration of Arid Land Soils.

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.

Strategies for restoring the structure and function of lichen‐moss biocrust communities

Biological soil crusts (biocrusts) are crucial components of dryland ecosystems, but they are slow to recover following disturbance. Herein, we evaluated several methods for restoring lichen‐moss biocrusts that included factorial applications of moss fragments in a water‐slurry (1) with and without lichen fragments (to restore biocrust taxonomic structure), (2) with and without clay (to facilitate establishment), and (3) with and without jute ground cloth (to facilitate establishment). Three and four years after inoculation, moss and lichen cover was up to five and eight times higher on jute ground cloth than on bare ground, respectively. Lichen cover was six times higher in plots where lichen fragments were added. Clay amendments did not increase moss or lichen establishment. To understand the effects of biocrust recovery on soil properties, we measured soil inorganic nitrogen, microbial biomass carbon, and soil water availability in restoration and control plots. Restored biocrusts decreased inorganic NH4‐N availability by 67% when compared to controls 3 years after inoculation, but did not influence the availability of inorganic NO3‐N, soil water, or microbial biomass carbon. Our results demonstrate that adding a biocrust inoculant to jute ground cloth can expedite recovery of lichen‐moss biocrust and reestablish its influence on soil properties within a few years.

Temporal and abiotic fluctuations may be preventing successful rehabilitation of soil‐stabilizing biocrust communities

Land degradation is a persistent ecological problem in many arid and semiarid systems globally (drylands hereafter). Most instances of dryland degradation include some form of soil disturbance and/or soil erosion, which can hinder vegetation establishment and reduce ecosystem productivity. To combat soil erosion, researchers have identified a need for rehabilitation of biological soil crusts (biocrusts), a globally relevant community of organisms aggregating the soil surface and building soil fertility. Here, the impact of plant and biocrust cover was tested on soil erosion potential in the piñon-juniper woodlands of Bandelier National Monument, New Mexico, USA. Biocrusts were found to be similarly influential to vascular plants in reducing erosion, largely acting by promoting surface roughness. The potential to rehabilitate biocrusts within the Monument was also tested. Plots were inoculated on eroding soils before the summer monsoon with greenhouse-cultured biocrusts. In a full-factorial design, treatments to reduce or halt erosion were administered to the inoculated plots and their paired controls. These erosion-reduction treatments included barriers to overland flow (flashing), slash placement, and seeding of vascular plants. Dynamic changes to soil stability, penetration resistance, and extractable soil nutrients were observed through time, but no strong effects with the addition of biocrust inoculum, seeding, or erosion intervention treatments were seen. The results do suggest possible ways forward to successfully rehabilitate biocrust, including varying the timing of biocrust application, amending inoculum application with different types of soil stabilization techniques, and adding nutrients to soils. The insights gleaned from the lack of response brings us closer to developing effective techniques to arrest soil loss in these socially and ecologically important dryland systems.

Soil characteristics are associated with gradients of big sagebrush canopy structure after disturbance

AbstractReestablishing shrub canopy cover after disturbance in semi‐arid ecosystems, such as sagebrush steppe, is essential to provide wildlife habitat and restore ecosystem functioning. While several studies have explored the effects of landscape and climate factors on the success or failure of sagebrush seeding, the influence of soil properties on gradients of shrub canopy structure in successfully seeded areas remains largely unexplored. In this study, we evaluated associations between soil properties and gradients in sagebrush canopy structure in stands that had successfully reestablished after fire and subsequent seeding treatments. Using a dataset collected across the Great Basin, USA, of sagebrush stands that had burned and reestablished between 1986 and 2013, we tested soil depth and texture, soil surface classification, biological soil crusts plus mean historical precipitation, solar heatload, and fire history as modeling variables to explore gradients in sagebrush canopy structure growth in terms of cover, height, and density. Deeper soils were associated with greater sagebrush canopy structure development in terms of plant density and percent cover, coarser textured soils were associated with greater sagebrush cover and density, and more clayey soils were typically associated with greater height. Biological crust presence was also positively associated with enhanced sagebrush canopy growth, but adding more demographically or morphologically explicit descriptions of biocrust communities did not improve explanatory power. Increasing heatload had a negative effect on sagebrush canopy structure growth, and increased mean annual precipitation was only associated with greater sagebrush height. Given that conservation and restoration of the sagebrush steppe ecosystems has become a priority for land managers, the associations we identify between gradients in post‐fire sagebrush canopy structure growth and field‐identifiable soil characteristics may improve planning of land treatments for sagebrush restoration and the understanding of semi‐arid ecosystem functioning and post‐disturbance dynamics.

Spatial segregation of the biological soil crust microbiome around its foundational cyanobacterium, Microcoleus vaginatus, and the formation of a nitrogen-fixing cyanosphere

BackgroundBiological soil crusts (biocrusts) are a key component of arid land ecosystems, where they render critical services such as soil surface stabilization and nutrient fertilization. The bundle-forming, filamentous, non-nitrogen-fixing cyanobacterium Microcoleus vaginatus is a pioneer primary producer, often the dominant member of the biocrust microbiome, and the main source of leaked organic carbon. We hypothesized that, by analogy to the rhizosphere of plant roots, M. vaginatus may shape the microbial populations of heterotrophs around it, forming a specialized cyanosphere.ResultsBy physically isolating bundles of M. vaginatus from biocrusts, we were able to study the composition of the microbial populations attached to it, in comparison to the bulk soil crust microbiome by means of high-throughput 16S rRNA sequencing. We did this in two M. vaginatus-dominated biocrust from distinct desert biomes. We found that a small, selected subset of OTUs was significantly enriched in close proximity to M. vaginatus. Furthermore, we also found that a majority of bacteria (corresponding to some two thirds of the reads) were significantly more abundant away from this cyanobacterium. Phylogenetic placements suggest that all typical members of the cyanosphere were copiotrophs and that many were diazotrophs (Additional file 1: Tables S2 and S3). Nitrogen fixation genes were in fact orders of magnitude more abundant in this cyanosphere than in the bulk biocrust soil as assessed by qPCR. By contrary, competition for light, CO2, and low organic carbon concentrations defined at least a part of the OTUs segregating from the cyanobacterium.ConclusionsWe showed that M. vaginatus acts as a significant spatial organizer of the biocrust microbiome. On the one hand, it possesses a compositionally differentiated cyanosphere that concentrates the nitrogen-fixing function. We propose that a mutualism based on C for N exchange between M. vaginatus and copiotrophic diazotrophs helps sustains this cyanosphere and that this consortium constitutes the true pioneer community enabling the colonization of nitrogen-poor soils. On the other hand, a large number of biocrust community members segregate away from the vicinity of M. vaginatus, potentially through competition for light or CO2, or because of a preference for oligotrophy.

Responses of biological soil crusts to rehabilitation strategies

Biological soil crusts (biocrusts) are common to dryland ecosystems and can influence a broad suite of soil ecological functions including stability and surface hydrology. Due to long recovery times following disturbance, there is a clear need for rehabilitation strategies to enhance the recovery of biocrust communities. Essential to biocrust recovery are exopolysaccharides (EPS): secretions comprised mainly of high molecular weight polymers that protect cyanobacteria and other biocrust organisms from harsh environmental conditions. We examined whether biocrust rehabilitation strategies (combinations of inoculation with surface shading and artificial soil stabilization) promote EPS production. To test if responses varied by soil texture, we measured biocrust recovery on two fine-textured soils (clay and sandy clay loam) in a cool desert ecosystem. Shade coupled with inoculum addition resulted in the highest biocrust recovery, especially on clay soils. Independent of rehabilitation strategies, natural recovery of biocrusts occurred more rapidly on clay soils, reflected by greater increases in chlorophyll a (chl a). Chl a, a proxy for cyanobacterial biomass, was correlated to EPS amounts, suggesting that cyanobacteria are significant contributors to EPS production in biocrust development. Despite the role of EPS in biocrust establishment, EPS amounts had negligible effects on soil stability on the fine soil texture.

Soil moisture dynamics under two rainfall frequency treatments drive early spring CO2 gas exchange of lichen-dominated biocrusts in central Spain.

BackgroundBiocrusts, communities dominated by mosses, lichens, cyanobacteria, and other microorganisms, largely affect the carbon cycle of drylands. As poikilohydric organisms, their activity time is often limited to short hydration events. The photosynthetic and respiratory response of biocrusts to hydration events is not only determined by the overall amount of available water, but also by the frequency and size of individual rainfall pulses.MethodsWe experimentally assessed the carbon exchange of a biocrust community dominated by the lichen Diploschistes diacapsis in central Spain. We compared the effect of two simulated precipitation patterns providing the same overall amount of water, but with different pulse sizes and frequency (high frequency: five mm/day vs. low frequency: 15 mm/3 days), on net/gross photosynthesis and dark respiration.ResultsRadiation and soil temperature, together with the watering treatment, affected the rates of net and gross photosynthesis, as well as dark respiration. On average, the low frequency treatment showed a 46% ± 3% (mean ± 1 SE) lower rate of net photosynthesis, a 13% ± 7% lower rate of dark respiration, and a 24% ± 8% lower rate of gross photosynthesis. However, on the days when samples of both treatments were watered, no differences between their carbon fluxes were observed. The carbon flux response of D. diacapsis was modulated by the environmental conditions and was particularly dependent on the antecedent soil moisture.DiscussionIn line with other studies, we found a synergetic effect of individual pulse size, frequency, environmental conditions, and antecedent moisture on the carbon exchange fluxes of biocrusts. However, most studies on this subject were conducted in summer and they obtained results different from ours, so we conclude that there is a need for long-term experiments of manipulated precipitation impacts on the carbon exchange of biocrusts. This will enable a more complete assessment of the impacts of climate change-induced alterations in precipitation patterns on biocrust communities.

The influence of climatic legacies on the distribution of dryland biocrust communities.

Predicting the distribution of biocrust species, mosses, lichens and liverworts associated with surface soils is difficult, but climatic legacies (changes in climate over the last 20k years) can improve our prediction of the distribution of biocrust species. To provide empirical support for this hypothesis, we used a combination of network analyses and structural equation modelling to identify the role of climatic legacies in predicting the distribution of ecological clusters formed by species of mosses, lichens and liverworts using data from 282 large sites distributed across 0.6 million km2 of eastern Australia. Two ecological clusters contained 87% of the 120 moss, lichen and liverwort species. Both clusters contained lichen, moss and liverwort species, but were dominated by different families. Sites where the air temperature increased the most over 20kyears (positive temperature legacies) were associated with reductions in the relative abundance of species from the lichen (Peltulaceae and Teloschistaceae) and moss (Bryaceae) families (Cluster A species), greater groundstorey plant cover and lower soil pH. Sites where precipitation has increased over the past 20kyears (positive precipitation legacy) were associated with increases in the relative abundance of lichen (Cladoniaceae, Lecideaceae and Thelotremataceae) and moss (Pottiaceae) families (Cluster B species) and lower levels of soil pH. Sites where temperatures have increased the most in the past 20kyears suppressed the negative effects of plant cover on Cluster B by reducing plant cover. Increased intensity of grazing suppressed the negative effect of soil pH and the positive effect of soil carbon, on the relative abundance of Cluster B taxa. Finally, increasing temperature and precipitation legacies reduced the negative effect of soil pH on Cluster B. Understanding of the importance of climatic legacies improves our ability to predict how biocrust assemblies might respond to ongoing global environmental change associated with increasing land use intensification, increasing temperature and reduced rainfall.

Insolation and disturbance history drive biocrust biodiversity in Western Montana rangelands

Background and AimsBiological soil crust (biocrust) communities, though common and important in the intermountain west, have received little research attention. There are gaps in understanding what influences biocrust species’ abundance and distributions in this ecoregion. Climatic, edaphic, topographic, and biotic forces, in addition to anthropogenic disturbance can all influence the biocrust.MethodsWe determined the relative influence of several possible environmental filters in biocrust communities of western Montana (USA) grasslands at two spatial scales. The larger scale exploited strong topographically-dictated climatic variation across >60km2, while the smaller scale focused on differences among distinct microsites within ~700m2 plots.ResultsWe detected a total of 96 biocrust taxa, mostly lichens. Biocrust richness at each site ranged from 0 to 39 species, averaging 14 species. Insolation, aspect, and disturbance history were the strongest predictors of biocrust richness, abundance, and species turnover across the landscape; soil texture was influential for some biocrust community properties. Steep, north-facing slopes that receive longer periods of shade harbored higher diversity and cover of biocrust than south-facing sites. At a small scale, interspaces among native herbaceous communities supported the greatest diversity of biocrust species, but microsites under shrub canopies supported the greatest cover.ConclusionsWe found that, among the variables investigated, tillage, insolation, soil texture and the associated vegetation community were the most important drivers of biocrust abundance and species richness. This study can inform the practice of restoration and conservation, and also guide future work to improve predictions of biocrust properties.

Cyanobacteria Inoculation Improves Soil Stability and Fertility on Different Textured Soils: Gaining Insights for Applicability in Soil Restoration

Cyanobacteria are ubiquitous components of biocrust communities and the first colonizers of terrestrial ecosystems. They play multiple roles in the soil by fixing C and N and synthesizing exopolysaccharides, which increase soil fertility and water retention and improve soil structure and stability. Application of cyanobacteria as inoculants to promote biocrust development has been proposed as a novel biotechnological technique for restoring barren degraded areas and combating desertification processes in arid lands. However, previous to their widespread application under field conditions, research is needed to ensure the selection of the most suitable species. In this study, we inoculated two cyanobacterial species, Phormidium ambiguum (non N-fixing) and Scytonema javanicum (N-fixing), on different textured soils (from silt loam to sandy), and analyzed cyanobacteria biocrust development and evolution of physicochemical soil properties for three months under laboratory conditions. Cyanobacteria inoculation led to biocrust formation in all soil types. Scanning electron microscope (SEM) images showed contrasting structure of the biocrust induced by the two cyanobacteria. The one from P. ambiguum was characterized by thin filaments that enveloped soil particles and created a dense, entangled network, while the one from S. javanicum consisted of thicker filaments that grouped as bunches in between soil particles. Biocrust development, assessed by chlorophyll a content and crust spectral properties, was higher in S. javanicum-inoculated soils compared to P. ambiguum-inoculated soils. Either cyanobacteria inoculation did not increase soil hydrophobicity. S. javanicum promoted a higher increase in total organic C and total N content, while P. ambiguum was more effective in increasing total exopolysaccharide (EPS) content and soil penetration resistance. The effects of cyanobacteria inoculation also differed among soil types and the highest improvement in soil fertility compared to non-inoculated soils was found in sandy and silty soils, which originally had lowest fertility. On the whole, the improvement in soil fertility and stability supports the viability of using cyanobacteria to restore degraded arid soils.

Fire and Grazing Influence Site Resistance to Bromus tectorum Through Their Effects on Shrub, Bunchgrass and Biocrust Communities in the Great Basin (USA)

Shrubs, bunchgrasses and biological soil crusts (biocrusts) are believed to contribute to site resistance to plant invasions in the presence of cattle grazing. Although fire is a concomitant disturbance with grazing, little is known regarding their combined impacts on invasion resistance. We are the first to date to test the idea that biotic communities mediate the effects of disturbance on site resistance. We assessed cover of Bromus tectorum, shrubs, native bunchgrasses, lichens and mosses in 99 burned and unburned plots located on similar soils where fires occurred between 12 and 23 years before sampling. Structural equation modeling was used to test hypothesized relationships between environmental and disturbance characteristics, the biotic community and resistance to B. tectorum cover. Characteristics of fire and grazing did not directly relate to cover of B. tectorum. Relationships were mediated through shrub, bunchgrass and biocrust communities. Increased site resistance following fire was associated with higher bunchgrass cover and recovery of bunchgrasses and mosses with time since fire. Evidence of grazing was more pronounced on burned sites and was positively correlated with the cover of B. tectorum, indicating an interaction between fire and grazing that decreases site resistance. Lichen cover showed a weak, negative relationship with cover of B. tectorum. Fire reduced near-term site resistance to B. tectorum on actively grazed rangelands. Independent of fire, grazing impacts resulted in reduced site resistance to B. tectorum, suggesting that grazing management that enhances plant and biocrust communities will also enhance site resistance.

Lichen-biocrust diversity in a fragmented dryland: Fine scale factors are better predictors than landscape structure

Biological soil crusts (or biocrusts) are widespread, diverse and important components of drylands sometimes threatened by global change drivers. However, their response to fragmentation processes is poorly known. The aim of this study was to assess the effects of changing landscape structure, given by land use change and the presence of linear infrastructure (e.g., roads), on the cover and diversity of lichen-biocrusts. We also evaluated the influence of several subrogates of fragment quality, such as soil properties, vascular plant community structure and topography. Biocrust cover and diversity were measured in 50 remnants of a Mediterranean shrubland. The fragments varied in size, connectivity and distance to a road, but also in plant and soil attributes, topography and fragment history. We applied general linear and mixed models to assess the effects of environmental variables on biocrust communities. Biocrust cover, richness and species composition were mostly unresponsive to changes in landscape structure, while connectivity and distance to the road decreased species diversity. Soil properties better explained the variation in biocrust cover and diversity. Changes in plant community and biocrust community composition were coupled. We also identified several biocrust species with strong capacity to reflect landscape structure. Our findings suggest that landscape structure needs to be evaluated jointly with other environmental factors to fully understand the consequences of fragmentation processes on biocrust communities and the subsequent implications for their functional role in drylands.

Species-specific nitrogenase activity in lichen-dominated biological soil crusts from the Colorado Plateau, USA

Background and aim Biological soil crusts (biocrusts) play numerous crucial roles in drylands, which comprise over 40% of Earth’s terrestrial surface. Among these key contributions is the fixation of atmospheric nitrogen. Yet, relatively little is known about the N2 fixation capabilities of different lichen species that are found in late successional biocrust communities across drylands globally.

Warming reduces the cover, richness and evenness of lichen-dominated biocrusts but promotes moss growth: insights from an 8yr experiment.

Despite the important role that biocrust communities play in maintaining ecosystem structure and functioning in drylands world-wide, few studies have evaluated how climate change will affect them. Using data from an 8-yr-old manipulative field experiment located in central Spain, we evaluated how warming, rainfall exclusion and their combination affected the dynamics of biocrust communities in areas that initially had low (<20%, LIBC plots) and high (>50%, HIBC plots) biocrust cover. Warming reduced the richness (35±6%), diversity (25±8%) and cover (82±5%) of biocrusts in HIBC plots. The presence and abundance of mosses increased with warming through time in these plots, although their growth rate was much lower than the rate of lichen death, resulting in a net loss of biocrust cover. On average, warming caused a decrease in the abundance (64±7%) and presence (38±24%) of species in the HIBC plots. Over time, lichens and mosses colonized the LIBC plots, but this process was hampered by warming in the case of lichens. The observed reductions in the cover and diversity of lichen-dominated biocrusts with warming will lessen the capacity of drylands such as that studied here to sequester atmospheric CO2 and to provide other key ecosystem services associated to these communities.

Biocrust carbon isotope signature was depleted under a C3 forb compared to interspace

Plants and biological soil crusts (biocrusts) are the key producers in drylands, but biocrusts seldom show net CO2 uptake. I hypothesized that biocrusts could augment CO2 fixation by incorporating plant-derived carbon. I collected biocrusts located at the base of Gutierrezia sarothrae (C3 forb), Bouteloua gracilis (C4 grass), and from bare interspaces between plants, and from a mesocosm experiment with live B. gracilis or dead B. gracilis roots. To trace carbon sources, I determined 13C values of the biocrust community, isolated cyanobacteria and lichen, and plant leaves because the photosynthetic pathway distinguishes the tissue 13C values. Biocrust communities and washed cyanobacteria and cyanolichen in G. sarothrae microsites were depleted by ~2‰ relative to other locations. Biocrust δ13C did not differ between the interspace and live or dead B. gracilis. Potential mechanisms for the trend in biocrust δ13C adjacent to C3 plants include differences in microsite conditions, biocrust communities, use of respired CO2 in the soil matrix for photosynthesis, or mixotrophic use of plant photosynthates. Further investigation of this observation may improve understanding of the degree to which the activities of dryland primary producers are coupled.

Diversity of biocrust-forming cyanobacteria in a semiarid gypsiferous site from Central Spain

Cyanobacteria are a key constituent of biocrusts, communities dominated by lichens, mosses and associated microorganisms, which are prevalent in drylands worldwide and that largely determine their functioning. Despite their importance, there are large gaps in our knowledge of the composition and diversity of cyanobacteria associated with biocrusts, particularly in areas such as the Mediterranean Basin. We studied the diversity of these cyanobacteria in a gypsiferous grassland from Central Spain using both morphological identification after cultivation and genetic analyses with the 16 S rRNA gene. Nine different morphotypes were observed, eight corresponding to filamentous, and one to unicellular cyanobacteria. We found cyanobacterial genera typical of biocrust communities, such as Microcoleus and Trichocoleus, and N-fixing cyanobacteria such as Scytonema and Nostoc. Genetic information allowed us to identify cultures belonging to recently described genera such as Roholtiella, Nodosilinea and Mojavia. We also describe two new phylotypes of Microcoleus and Scytonema, which are key genera contributing to ecosystem functioning in biocrust-dominated ecosystems worldwide.