What is a biocrust? A refined, contemporary definition for a broadening research community.

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.

Incorporating Biogeochemistry into Dryland Restoration.

Dryland degradation is a persistent and accelerating global problem. Although the mechanisms initiating and maintaining dryland degradation are largely understood, returning productivity and function through ecological restoration remains difficult. Water limitation commonly drives slow recovery rates within drylands; however, the altered biogeochemical cycles that accompany degradation also play key roles in limiting restoration outcomes. Addressing biogeochemical changes and resource limitations may help improve restoration efforts within this difficult-to-restore biome. In the present article, we present a synthesis of restoration literature that identifies multiple ways biogeochemical understandings might augment dryland restoration outcomes, including timing restoration around resource cycling and uptake, connecting heterogeneous landscapes, manipulating resource pools, and using organismal functional traits to a restoration advantage. We conclude by suggesting ways to incorporate biogeochemistry into existing restoration frameworks and discuss research directions that may help improve restoration outcomes in the world's highly altered dryland landscapes.

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.

Spectrally monitoring the response of the biocrust moss Syntrichia caninervis to altered precipitation regimes.

Climate change is expected to impact drylands worldwide by increasing temperatures and changing precipitation patterns. These effects have known feedbacks to the functional roles of dryland biological soil crust communities (biocrusts), which are expected to undergo significant climate-induced changes in community structure and function. Nevertheless, our ability to monitor the status and physiology of biocrusts with remote sensing is limited due to the heterogeneous nature of dryland landscapes and the desiccation tolerance of biocrusts, which leaves them frequently photosynthetically inactive and difficult to assess. To address this critical limitation, we subjected a dominant biocrust species Syntrichia caninervis to climate-induced stress in the form of small, frequent watering events, and spectrally monitored the dry mosses' progression towards mortality. We found points of spectral sensitivity responding to experimentally-induced stress in desiccated mosses, indicating that spectral imaging is an effective tool to monitor photosynthetically inactive biocrusts. Comparing the Normalized Difference Vegetation Index (NDVI), the Simple Ratio (SR), and the Normalized Pigment Chlorophyll Index (NPCI), we found NDVI minimally effective at capturing stress in precipitation-stressed dry mosses, while the SR and NPCI were highly effective. Our results suggest the strong potential for utilizing spectroscopy and chlorophyll-derived indices to monitor biocrust ecophysiological status, even when biocrusts are dry, with important implications for improving our understanding of dryland functioning.