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Vegetation and precipitation shifts interact to alter organic and inorganic carbon storage in cold desert soils

Dryland ecosystems are experiencing shifts in rainfall and plant community composition, which are expected to alter cycling and storage of soil carbon (C). Few experiments have been conducted to examine long‐term effects on (1) soil organic C (SOC) pools throughout the soil profile, and (2) soil inorganic C (SIC) pools as they relate to dynamic changes in C storage and climate change. We measured SOC and SIC from 0 to 1 m beneath plants and in adjacent interplant microsites following nearly 20 yr of experimental manipulations of plant community (native sagebrush steppe or monoculture of exotic crested wheatgrass) and the amount and timing of water availability (ambient, or doubling of annual rainfall in the dormant, DORM, or growing, GROW, season). Under sagebrush plants, GROW increased both SOC and SIC pools, resulting in total carbon (TC) pools 15% greater than plots receiving ambient precipitation, while DORM decreased SOC and SIC pools, decreasing TC pools 20% from ambient. Under crested wheatgrass plants, GROW increased SOC by 73% but decreased SIC by 11% relative to ambient, netting no change in TC pools, while DORM SIC pools were 5% greater than ambient, with no significant increase in either SOC or TC pools. GROW significantly increased TC pools for interplant microsites, regardless of vegetation treatment. At the community scale and summing C pools weighted by percent patch cover, patterns of TC pool were similar to plot measurements. Our findings suggest that sagebrush communities can become a net C source to the atmosphere with increases in dormant season rainfall rather than a C sink as previously predicted . We also provide evidence of SIC as an important and dynamic C sequestration mechanism in drylands. Consideration of vegetation type, all or most of the soil profile, and both organic and inorganic C pools are all important to accurately predict C sequestration with changing climate and disturbance in drylands.

Ecosphere

Woody plant encroachment of grassland and the reversibility of shrub dominance: Erosion, fire, and feedback processes

Many grass-dominated ecosystems in dryland regions have experienced increasing woody plant density and abundance during the past century. In many cases, this process has led to land degradation and declines in ecosystem functions. An example is the Chihuahuan Desert in the southwestern United States, which experienced different stages of shrub encroachment in the past 150 years. Among a wide variety of mechanisms to explain the grass–shrub transitions in this dryland system, soil erosion (both wind and water) and fire are particularly well studied. Here, we synthesize recent developments on the drivers and feedback in the process of shrub encroachment in the Chihuahuan Desert through the intercomparison of two Long Term Ecological Research (LTER) sites, namely Jornada and Sevilleta. Experimental and modeling studies support a conceptual framework, which underscores the important roles of erosion and fire in woody plant encroachment. Collectively, research at the Jornada LTER provided complementary, quantitative support to the well-known fertile-islands framework. Studies at the Sevilleta LTER expanded the framework, adding fire as a major disturbance to woody plants. Conceptual models derived from the synthesis represent the general understanding of shrub encroachment that emerged from research at these two sites, and can guide management interventions aimed at reducing or mitigating undesirable ecosystem state change in many other drylands worldwide.

New Mexico

Soil texture and precipitation seasonality influence plant community structure in North American temperate shrub steppe

In drylands, the coexistence of grasses and woody plants has been attributed to soil‐water resource partitioning. Soil texture and precipitation seasonality can influence the amount and distribution of water in the soil, and their interaction may play an important role in determining the relative importance of grasses and woody plants. We investigated the influence of this interaction on plant functional types across a broad range of precipitation regimes and soil textures in western North America by analyzing plant‐cover data collected at 2,084 plots that included the widespread shrub big sagebrush ( Artemisia tridentata Nutt.). We characterized how the significance of the inverse‐texture effect varies across soil conditions by quantifying relationships between precipitation and foliar cover on finer‐ vs. coarser‐textured soils across a range of potential texture divisions represented by sand content. We found evidence of the inverse‐texture effect for every plant functional type (except for cheatgrass) that we examined with at least one component of precipitation (annual, warm, or cold season), and provide the first evidence for this effect in locations with cold‐season‐dominated precipitation regimes. The texture and precipitation combinations that exhibited the inverse‐texture effect varied with plant functional type, presumably because of effects of soil texture on water availability at different soil depths with season. Furthermore, we found an inverse‐texture effect that was remarkably similar for shrub cover with cold‐season precipitation and grass cover with warm‐season precipitation. These results provide new insight into how the inverse‐texture effect interacts with precipitation seasonality to influence plant functional type composition in drylands, and further suggest that quantifying the soil‐texture division at which the inverse‐texture effect is relevant under a given set of environmental conditions may provide support for the effect across dryland plant communities.

Arizona, California, Colorado, Idaho, Montana, New

Relations between rainfall–runoff-induced erosion and aeolian deposition at archaeological sites in a semi-arid dam-controlled river corridor

Process dynamics in fluvial-based dryland environments are highly complex with fluvial, aeolian, and alluvial processes all contributing to landscape change. When anthropogenic activities such as dam-building affect fluvial processes, the complexity in local response can be further increased by flood- and sediment-limiting flows. Understanding these complexities is key to predicting landscape behavior in drylands and has important scientific and management implications, including for studies related to paleoclimatology, landscape ecology evolution, and archaeological site context and preservation. Here we use multi-temporal LiDAR surveys, local weather data, and geomorphological observations to identify trends in site change throughout the 446-km-long semi-arid Colorado River corridor in Grand Canyon, Arizona, USA, where archaeological site degradation related to the effects of upstream dam operation is a concern. Using several site case studies, we show the range of landscape responses that might be expected from concomitant occurrence of dam-controlled fluvial sand bar deposition, aeolian sand transport, and rainfall-induced erosion. Empirical rainfall-erosion threshold analyses coupled with a numerical rainfall–runoff–soil erosion model indicate that infiltration-excess overland flow and gullying govern large-scale (centimeter- to decimeter-scale) landscape changes, but that aeolian deposition can in some cases mitigate gully erosion. Whereas threshold analyses identify the normalized rainfall intensity (defined as the ratio of rainfall intensity to hydraulic conductivity) as the primary factor governing hydrologic-driven erosion, assessment of false positives and false negatives in the dataset highlight topographic slope as the next most important parameter governing site response. Analysis of 4+ years of high resolution (four-minute) weather data and 75+ years of low resolution (daily) climate records indicates that dryland erosion is dependent on short-term, storm-driven rainfall intensity rather than cumulative rainfall, and that erosion can occur outside of wet seasons and even wet years. These results can apply to other similar semi-arid landscapes where process complexity may not be fully understood.

Arizona

Biological soil crusts: Diminutive communities of potential global importance

Biological soil crusts (biocrusts) are widespread, diverse communities of cyanobacteria, fungi, lichens, and mosses living on soil surfaces, primarily in drylands. Biocrusts can locally govern primary production, soil fertility, hydrology, and surface energy balance, with considerable variation in these functions across alternate community states. Further, these communities have been implicated in Earth system functioning via potential influences on global biogeochemistry and climate. Biocrusts are easily destroyed by disturbances and appear to be exceptionally vulnerable to warming temperatures and altered precipitation inputs, signaling possible losses of dryland functions with global change. Despite these concerns, we lack sufficient spatiotemporal data on biocrust function, cover, and community structure to confidently assess their ecological roles across the extensive dryland biome. Here, we present the case for cross-scale research and restoration efforts coupled with remote-sensing and modeling approaches that improve our collective understanding of biocrust responses to global change and the ecological roles of these diminutive communities at global scales.

Frontiers in Ecology and the Environment

Patterns and controls on nitrogen cycling of biological soil crusts

Biocrusts play a significant role in the nitrogen [N ] cycle within arid and semi-arid ecosystems, as they contribute major N inputs via biological fixation and dust capture, harbor internal N transformation processes, and direct N losses via N dissolved, gaseous and erosional loss processes (Fig. 1). Because soil N availability in arid and semi-arid ecosystems is generally low and may limit net primary production (NPP), especially during periods when adequate water is available, understanding the mechanisms and controls of N input and loss pathways in biocrusts is critically important to our broader understanding of N cycling in dryland environments. In particular, N cycling by biocrusts likely regulates short-term soil N availability to support vascular plant growth, as well as long-term N accumulation and maintenance of soil fertility. In this chapter, we review the influence of biocrust nutrient input, internal cycling, and loss pathways across a range of biomes. We examine linkages between N fixation capabilities of biocrust organisms and spatio-temporal patterns of soil N availability that may influence the longer-term productivity of dryland ecosystems. Lastly, biocrust influence on N loss pathways such as N gas loss, leakage of N compounds from biocrusts, and transfer in wind and water erosion are important to understand the maintenance of dryland soil fertility over longer time scales. Although great strides have been made in understanding the influence of biocrusts on ecosystem N cycling, there are important knowledge gaps in our understanding of the influence of biocrusts on ecosystem N cycling that should be the focus of future studies. Because work on the interaction of N cycling and biocrusts was reviewed in Belnap and Lange (2003), this chapter will focus primarily on research findings that have emerged over the last 15 years (2000-2015).

Book chapter

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 N 2 fixation capabilities of different lichen species that are found in late successional biocrust communities across drylands globally. Methods In order to improve our species-specific understanding of biocrust lichen N 2 fixation, we collected biocrusts dominated by four common species of lichens – Collema spp., Gyalolechia desertorum , Psora decipiens , and Squamarina lentigera – that represent a range of lichen families and morphological types. Nitrogenase activity of the biocrust community dominated by these lichens was evaluated using the acetylene reduction assay. Additionally, biocrust community composition was assessed using the point-intercept method along transects at varied distances from exposed bedrock. Results As expected, Collema spp.-dominated biocrusts had the highest rates of nitrogenase activity, with rates up to seven times larger than those of the other three target species. Nitrogen concentrations and carbon:nitrogen ratios of lichen tissue differed among lichen species. However, when the composite biocrust profile was assessed (i.e., biocrust tissue, microbial cells, and mineral soil to a 2 cm depth) these among-species differences in total nitrogen disappeared. Community composition changed according to distance from exposed bedrock, with a higher diversity of lichens closer to the bedrock. Conclusion Multiple drivers, including climate and land use change, affect biocrust community composition and species-specific functional information, even within a group such as late successional biocrusts, could help in forecasting the potential effects of global change on N 2 fixation, and consequently, soil fertility in drylands.

Colorado Plateau

Understanding the links between functional performance and environment in cold desert ecosystems through the flagship biocrust forming moss Syntrichia caninervis

Background and aims Biocrust communities shape the soil surface in drylands, regulating important ecosystem properties. Despite this, few works address key functional attributes in these poikilohydric photosynthetic communities, including the length of metabolic activity in the field and its relationship with microclimate. We aim to disentangle the links between functional performance of a keystone biocrust species and environment in drylands, in order to better understand ecosystem functioning and services provided to the soil by these organisms. Methods We developed a 23 months monitoring experiment using very high resolution measurements of chlorophyll a fluorescence and microclimate in the field, comparing habitats with soils of different particle size composition and microclimatic conditions for the flagship biocrust moss Syntrichia caninervis on the Colorado Plateau, Utah, USA. Results While growing in soils of different composition did not impair the functional performance of the species, the sun/shade plasticity did affect key physiological attributes. Our results quantified a strategy based on avoidance of the most extreme environmental conditions, where metabolic activity only occur during 19.9% of the total time monitored. Relative humidity was the best microclimatic predictor of metabolic activity, and the annual length of metabolically active periods showed strong convergences with other drylands of the world. Conclusions These results collectively suggest a strong control of the environment over the ecosystem services provided by biocrusts. Our results include evidence for an improved understanding of the plasticity of S. caninervis to its variety of habitats, and its response to potential threats under different climate change scenarios.

Utah

Shrubland carbon sink depends upon winter water availability in the warm deserts of North America

Global-scale studies suggest that dryland ecosystems dominate an increasing trend in the magnitude and interannual variability of the land CO 2 sink. However, such model-based analyses are poorly constrained by measured CO 2 exchange in open shrublands, which is the most common global land cover type, covering ∼14% of Earth’s surface. Here we evaluate how the amount and seasonal timing of water availability regulate CO 2 exchange between shrublands and the atmosphere. We use eddy covariance data from six US sites across the three warm deserts of North America with observed ranges in annual precipitation of ∼100–400mm, annual temperatures of 13–18°C, and records of 2–8 years (33 site-years in total). The Chihuahuan, Sonoran and Mojave Deserts present gradients in both mean annual precipitation and its seasonal distribution between the wet-winter Mojave Desert and the wet-summer Chihuahuan Desert. We found that due to hydrologic losses during the wettest summers in the Sonoran and Chihuahuan Deserts, evapotranspiration (ET) was a better metric than precipitation of water available to drive dryland CO 2 exchange. In contrast with recent synthesis studies across diverse dryland biomes, we found that NEP could not be directly predicted from ET due to wintertime decoupling of the relationship between ecosystem respiration (R eco ) and gross ecosystem productivity (GEP). Ecosystem water use efficiency (WUE=GEP/ET) did not differ between winter and summer. Carbon use efficiency (CUE=NEP/GEP), however, was greater in winter because R eco returned a smaller fraction of carbon to the atmosphere (23% of GEP) than in summer (77%). Combining the water-carbon relations found here with historical precipitation since 1980, we estimate that lower average winter precipitation during the 21st century reduced the net carbon sink of the three deserts by an average of 6.8TgC yr 1 . Our results highlight that winter precipitation is critical to the annual carbon balance of these warm desert shrublands.

Agricultural and Forest Meteorology

Vertical movement of soluble carbon and nutrients from biocrusts to subsurface mineral soils

Dryland ecosystems can be constrained by low soil fertility. Within drylands, the soil nutrient and organic carbon (C) cycling that does occur is often mediated by soil surface communities known as biological soil crusts (biocrusts), which cycle C and nutrients in the top ca. 0–2 cm of soil. However, the degree to which biocrusts are influencing soil fertility and biogeochemical cycling in deeper, subsurface mineral soils is unclear. The movement of dissolved resources from biocrusts to deeper soil layers in leachate may be one of the main mechanisms through which biocrust fertility is transferred downward towards deeper microbial communities and plant roots occurring within mineral soil. Here we examined the role of biocrust leachate in contributing to subsurface nutrient and soluble C pools and subsurface microbial cycling. We collected biocrusts from three biocrust successional stages and explored resource pools in situ at multiple soil depths, while collecting leachate and measuring nutrient and organic C concentrations and metabolite composition from each successional stage in the laboratory. After four leachate collections, we conducted an incubation of mineral soil collected from below each biocrust successional stage to measure heterotrophic microbial CO 2 flux and biomass. Overall, our findings observed that the degree of nutrient and C connectivity between biocrusts and the sub-crust mineral soil depended on the biocrust successional stage and the element being considered, and the influence of biocrust successional stage on mineral soil CO 2 flux is likely related to long-term resource build up. Together, our results suggest that the influence of biocrust leachate on subsurface mineral soil is complex and context dependent, but, over longer time periods and at later successional stages, can have measurable effects on dryland soil biogeochemical cycling with feedbacks to resource availability and CO 2 flux.

Geoderma

Effects of river regulation on aeolian landscapes, Colorado River, southwestern USA

Connectivity between fluvial and aeolian sedimentary systems plays an important role in the physical and biological environment of dryland regions. This study examines the coupling between fluvial sand deposits and aeolian dune fields in bedrock canyons of the arid to semiarid Colorado River corridor, southwestern USA. By quantifying significant differences between aeolian landscapes with and without modern fluvial sediment sources, this work demonstrates for the first time that the flow- and sediment-limiting effects of dam operations affect sedimentary processes and ecosystems in aeolian landscapes above the fluvial high water line. Dune fields decoupled from fluvial sand supply have more ground cover (biologic crust and vegetation) and less aeolian sand transport than do dune fields that remain coupled to modern fluvial sand supply. The proportion of active aeolian sand area also is substantially lower in a heavily regulated river reach (Marble–Grand Canyon, Arizona) than in a much less regulated reach with otherwise similar environmental conditions (Cataract Canyon, Utah). The interconnections shown here among river flow and sediment, aeolian sand transport, and biologic communities in aeolian dunes demonstrate a newly recognized means by which anthropogenic influence alters dryland environments. Because fluvial–aeolian coupling is common globally, it is likely that similar sediment-transport connectivity and interaction with upland ecosystems are important in other dryland regions to a greater degree than has been recognized previously.

Colorado

Effects of native forest restoration on soil hydraulic properties, Auwahi, Maui, Hawaiian Islands

Over historic time Hawai'i's dryland forests have been largely replaced by grasslands for grazing livestock. On-going efforts have been undertaken to restore dryland forests to bring back native species and reduce erosion. The reestablishment of native ecosystems on land severely degraded by long-term alternative use requires reversal of the impacts of erosion, organic-matter loss, and soil structural damage on soil hydraulic properties. This issue is perhaps especially critical in dryland forests where the soil must facilitate native plants' optimal use of limited water. These reforestation efforts depend on restoring soil ecological function, including soil hydraulic properties. We hypothesized that reforestation can measurably change soil hydraulic properties over restoration timescales. At a site on the island of Maui (Hawai'i, USA), we measured infiltration capacity, hydrophobicity, and abundance of preferential flow channels in a deforested grassland and in an adjacent area where active reforestation has been going on for fourteen years. Compared to the nearby deforested rangeland, mean field-saturated hydraulic conductivity in the newly restored forest measured by 55 infiltrometer tests was greater by a factor of 2.0. Hydrophobicity on an 8-point scale increased from average category 6.0 to 6.9. A 4-point empirical categorization of preferentiality in subsurface wetting patterns increased from an average 1.3 in grasslands to 2.6 in the restored forest. All of these changes act to distribute infiltrated water faster and deeper, as appropriate for native plant needs. This study indicates that vegetation restoration can lead to ecohydrologically important changes in soil hydraulic properties over decadal time scales.

Hawai'i

Unforeseen plant phenotypic diversity in a dry and grazed world

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 , 5 , 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 , 9 , 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.

Nature

Shrub influence on soil carbon and nitrogen in a semi-arid grassland is mediated by precipitation and largely insensitive to livestock grazing

Dryland (arid and semi-arid) ecosystems globally provide more than half of livestock production and store roughly one-third of soil organic carbon (SOC). Biogeochemical pools are changing due to shrub encroachment, livestock grazing, and climate change. We assessed how vegetation microsite, grazing, and precipitation interacted to affect SOC and total nitrogen (TN) at a site with long-term grazing manipulations and well-described patterns of shrub encroachment across elevation and mean annual precipitation (MAP) gradients. We analyzed SOC and TN in the context of vegetation cover at ungrazed locations within livestock exclosures, high-intensity grazing locations near water sources, and moderate-intensity grazing locations away from water. SOC was enhanced by MAP ( p < 0.0001), but grazing intensity had little effect regardless of MAP ( p = 0.12). Shrubs enhanced SOC (300–1279 g C m −2 ) and TN (27–122 g N m −2 ), except at high MAP where the contribution or stabilization of shrub inputs relative to grassland inputs was likely diminished. Cover of perennial herbaceous plants and litter were significant predictors of SOC ( r 2 = 0.63 and 0.34, respectively) and TN ( r 2 = 0.64 and 0.30, respectively). Our results suggest that continued shrub encroachment in drylands can increase SOC storage when grass production remains high, although this response may saturate with higher MAP. In contrast, grazing – at least at the intensities of our sites – has a lesser effect. These effects underscore the need to understand how future climate and grazing may interact to influence dryland biogeochemical cycling.

Arid Land Research and Management

Bacterial, fungal, and plant communities exhibit no biomass or compositional response to two years of simulated nitrogen deposition in a semiarid grassland

Nitrogen (N) deposition affects myriad aspects of terrestrial ecosystem structure and function, and microbial communities may be particularly sensitive to anthropogenic N inputs. However, our understanding of N deposition effects on microbial communities is far from complete, especially for drylands where data are comparatively rare. To address the need for an improved understanding of dryland biological responses to N deposition, we conducted a two-year fertilization experiment in a semiarid grassland on the Colorado Plateau in the southwestern United States. We evaluated effects of varied levels of N inputs on archaeal, bacterial, fungal and chlorophyte community composition within three microhabitats: biological soil crusts (biocrusts), soil below biocrusts, and the plant rhizosphere. Surprisingly, N addition did not affect the community composition or diversity of any of these microbial groups; however, microbial community composition varied significantly among sampling microhabitats. Further, while plant richness, diversity, and cover showed no response to N addition, there were strong linkages between plant properties and microbial community structure. Overall, these findings highlight the potential for some dryland communities to have limited biotic ability to retain augmented N inputs, possibly leading to large N losses to the atmosphere and to aquatic systems.

Environmental Microbiology

Biocrust mosses and cyanobacteria exhibit distinct carbon uptake responses to variations in precipitation amount and frequency

Dryland organisms exhibit varied responses to changes in precipitation, including event size, frequency, and soil moisture duration, influencing carbon uptake and reserve management strategies. This principle, central to the pulse-reserve paradigm, has not been thoroughly evaluated in biological soil crusts (biocrusts), essential primary producers on dryland surfaces. We conducted two experiments to investigate carbon uptake in biocrusts under different precipitation regimes. In the first, we applied a gradient of watering amounts to biocrusts dominated by moss or cyanobacteria, hypothesising distinct pulse-response strategies. The second experiment extended watering treatments over three months, varying pulse size and frequency. Our results revealed distinct carbon uptake patterns: moss crusts exhibited increased CO2 uptake with larger, less frequent watering events, whereas cyanobacteria crusts maintained similar carbon uptake across all event sizes. These findings suggest divergent pulse-response strategies across biocrust types, with implications for modelling dryland carbon dynamics and informing land management under changing precipitation regimes.

Utah

Droughting a megadrought: Ecological consequences of a decade of experimental drought atop aridification on the Colorado Plateau

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 (C 3 and C 4 grasses and C 3 and C 4 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.

Global Change Biology

Biocrust ecology: Unifying micro- and macro-scales to confront global change

Biological soil crusts (biocrusts) are communities of microbes, lichens and bryophytes living at the soil surface in drylands (Fig. 1; Belnap et al ., 2016). Biocrusts occur on all continents and can comprise a majority of cover in some systems (Belnap et al ., 2016). While species diversity and distributions have long been a research focus, interest in controls on community composition and cover has expanded as biocrusts are increasingly recognized for their roles in ecosystem functioning (Deane-Coe and Stanton, 2017). For example, biocrust organisms can stabilize soils (Belnap et al ., 2016; Faist et al ., 2017), fix atmospheric carbon (C) (Sancho et al ., 2016), and serve as the foremost source of ‘new’ soil nitrogen (N) in drylands, via N 2 fixation (Barger et al ., 2016) These contributions to gross primary production and soil fertility could be quite large, as high-end estimates suggest biocrusts and similar communities of bryophytes and lichens might account for 10% of terrestrial C- and 50% of N-fixation globally (Elbert et al ., 2012). Yet verifying these and other biocrust roles in ecosystem functioning is complicated by limited knowledge of biocrust cover and composition across the vast dryland biome (Ferrenberg et al ., 2017). It was against this backdrop that ‘Biocrust3: the 3 rd International workshop on biological soil crusts’ was held in Moab, UT, USA, on 26-30 September 2016. The workshop brought together over 50 scientists from 21 countries and six continents, and included numerous biocrust science pioneers (Fig. 2). The meeting was notable for its cross-scale focus, discussion of novel molecular and imaging techniques, and sessions on mapping and restoring biocrusts in a changing world. Here, we synthesize a central theme that emerged from Biocrust3, namely the potential for combining cutting edge tools with studies focused on organismal traits, ecosystem functions, and global change biology to advance the frontier of biocrust ecology.

New Phytologist