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the absorption and retention of moisture in the soil. In arid and semi-arid environments biocrust can cover over 70% of the soil not being covered by plants, indicating that the relationship between soil, water, and biocrust is extremely pertinent to these environments. Biocrusts has been shown to increase infiltration of water and pore space (or porosity) in soil but the opposite may occur depending on the type of biocrust. The effect biocrust has on water infiltration and the amount of water retained in the soil is greatly dependent on which microorganisms are most dominant in the specific forms of biocrust. Most research studies like that done by Canton et al. support that biological soil crust composed of large amounts of moss and lichens are better able to absorb water resulting in good soil infiltration. In comparison, biocrusts that aredominated by cyanobacteria is more likely to cause biological clogging where the pores of the soil are obstructed by the cyanobacteria responding to the presence of moisture by awakening from their dormancy and swelling. The darkening of the soil surface by biocrust can also raise the soil temperature leading to faster water evaporation. There is limited research, however, that indicates that the rough surface of cyanobacteria traps water runoff and lichen in cyanobacteria-dominant biocrust increase the porosity of the soil allowing for better infiltration than soil that does not have any biocrust.
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794:. Because crusts are only active when wet, some of these new conditions may reduce the amount of time when conditions are favorable for activity. Biological soil crusts require stored carbon when reactivating after being dry. If they do not have enough moisture to photosynthesize to make up for the carbon used, they can gradually deplete carbon stocks and die. Reduced carbon fixation also leads to decreased nitrogen fixation rates because crust organisms do not have sufficient energy for this energy-intensive process. Without carbon and nitrogen available, they are not able to grow nor repair damaged cells from excess radiation.
580:(a measure of the amount of light reflected off of the surface) compared to nearby soils, which increases the energy absorbed by the soil surface. Soils with well-developed biological soil crusts can be over 12 °C (22 °F) warmer than adjacent surfaces. Increased soil temperatures are associated with increased metabolic processes such as photosynthesis and nitrogen fixation, as well as higher soil water evaporation rates and delayed seedling germination and establishment. The activity levels of many arthropods and small mammals are also controlled by soil surface temperature.
692:. The increased micro-topography generally increases the probability that plant seeds will be caught on the soil surface and not blown away. Differences in water infiltration and soil moisture also contribute to differential germination depending on the plant species. It has been shown that while some native desert plant species have seeds with self-burial mechanisms that can establish readily in crusted areas, many exotic invasive plants do not. Therefore, the presence of biological soil crusts may slow the establishment of
626:. Cyanobacteria have evolved the ability to navigate the extreme conditions of their surrounding environment by existing in a biocrust. A trait of the biological soil crust community is that it will activate from a dormant state when it is exposed to precipitation transforming from a dry, dead-looking crust to an actively photosynthetic community. It will change its appearance to be vibrant and alive to the naked eye. Many crusts will even turn different shades of dark green. The cyanobacterium
776:, and tank treads can remove crusts and these disturbances have occurred over large areas globally. Once biological soil crusts are disrupted, wind and water can move sediments onto adjacent intact crusts, burying them and preventing photosynthesis of non-motile organisms such as mosses, lichens, green algae, and small cyanobacteria, and of motile cyanobacteria when the soil remains dry. This kills remaining intact crust and causes large areas of loss.
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401:. Frequently, single-celled organisms such as cyanobacteria or spores of free-living fungi colonize bare ground first. Once filaments have stabilized the soil, lichens and mosses can colonize. Appressed lichens are generally earlier colonizers or persist in more stressful conditions, while more three-dimensional lichens require long disturbance-free growth periods and more moderate conditions.
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depends on the dominant crust organisms, soil characteristics, and climate. In areas where biological soil crusts produce rough surface microtopography, water is detained longer on the soil surface and this increases water infiltration. However, in warm deserts where biological soil crusts are smooth
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The type of soil and its texture is also a major determining factor in the biological soil crust's relationship with water retention and filtration. Soils with a large presence of sand (less soil and clay) have high levels of water retention in their surface levels but have limited downward movement
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The amount of time it takes for the greening process in biocrust to occur varies on the environmental conditions in which the biocrust lives. Biocrust can take anywhere from five minutes to 24 hours to awaken from dormancy. The crusts will only awaken if the conditions are conducive to the biocrust.
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The morphology of biological soil crust surfaces can range from smooth and a few millimeters in thickness to pinnacles up to 15 cm high. Smooth biological soil crusts occur in hot deserts where the soil does not freeze, and consist mostly of cyanobacteria, algae, and fungi. Thicker and rougher
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and physical appearance of biological soil crusts vary depending on the climate, soil, and disturbance conditions. For example, biological soil crusts are more dominated by green algae on more acidic and less salty soils, whereas cyanobacteria are more favored on alkaline and haline soils. Within a
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Biocrust influences a soil's microtopography, carbohydrate content, porosity, and hydrophobicity which are the major contributing factors to soil hydrology. The relationship between biocrust and soil hydrology is not fully understood by scientists. It is known that the biocrust does play a role in
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and does not have the ability to maintain or regulate its own water retention. This causes the biocrust's water content to change depending on the water in the surrounding environment. Due to biological soil crust existing in mostly arid and semi-arid environments with the inability to hold water,
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or protection from disturbance are the easiest ways to maintain and improve biological soil crusts. Protection of relic sites that have not been disturbed can serve as reference conditions for restoration. There are several successful methods for stabilizing soil to allow recolonization of crusts
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of crust microorganisms which are active only when wet. Respiration can begin in as little as 3 minutes after wetting whereas photosynthesis reaches full activity after 30 minutes. Some groups have different responses to high water content, with some lichens showing decreased photosynthesis when
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Biological soil crusts cover about 12% of the earth's landmass. They are found on almost all soil types, but are more commonly found in arid regions of the world where plant cover is low and plants are more widely spaced. This is because crust organisms have a limited ability to grow upwards and
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Recovery following disturbance varies. Cyanobacteria cover can recover by propagules blowing in from adjacent undisturbed areas rapidly after disturbance. Total recovery of cover and composition occurs more rapidly in fine soil textured, moister environments (~2 years) and more slowly (>3800
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is able to return to a dormant state, migrating back down into the crust and bringing the pigment with it. This process goes along with the rapid turning on of metabolic pathways allowing metabolic functions to occur within the cells in the short periods of time when the crust is hydrated and
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Invasive species introduced by humans can also affect biological soil crusts. Invasive annual grasses can occupy areas once occupied by crusts and allow fire to travel between large plants, whereas previously it would have just jumped from plant to plant and not directly affected the crusts.
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migrates upward to the surface of the crust when hydrated, to perform oxygenic photosynthesis. In this photosynthetic process, the cyanobacteria carries with it a green-blue photosynthetic pigment to the surface of the crust. When inevitably there is a period of insufficient water again, the
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species and thus is less useful. Other methods such as fertilization and inoculation with material from adjacent sites may enhance crust recovery, but more research is needed to determine the local costs of disturbance. Today, direct inoculation of soil native microorganisms, bacteria and
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is one of the most dominant organisms found in biocrust and is fundamental to the crust's ability to reawaken from dormancy when rehydrated due to precipitation or runoff. Cyanobacteria have been found to outcompete the other components of biocrust when exposed to light and precipitation.
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Rajeev, Lara; da Rocha, Ulisses Nunes; Klitgord, Niels; Luning, Eric G.; Fortney, Julian; Axen, Seth D.; Shih, Patrick M.; Bouskill, Nicholas J.; Bowen, Benjamin P.; Kerfeld, Cheryl A.; Garcia-Pichel, Ferran; Brodie, Eoin L.; Northen, Trent R.; Mukhopadhyay, Aindrila (November 2013).
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requires energy from photosynthesis products, and thus increase with temperature given sufficient moisture. Nitrogen fixed by crusts has been shown to leak into surrounding substrate and can be taken up by plants, bacteria, and fungi. Nitrogen fixation has been recorded at rates of
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the crust is mainly dormant except for short periods of activity when the crust receives precipitation. Microorganisms like those that make up biological soil crust are good at responding quickly to changes in the environment even after a period of dormancy such as precipitation.
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Cyanobacteria are the main photosynthetic component of biological soil crusts, in addition to other photosynthetic taxa such as mosses, lichens, and green algae. The most common cyanobacteria found in soil crusts belong to large filamentous species such as those in the genus
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Biological soil crusts are extremely susceptible to disturbance from human activities. Compressional and shear forces can disrupt biological soil crusts especially when they are dry, leaving them to be blown or washed away. Thus, animal hoof impact, human footsteps,
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Desiccation can lead to oxidation and the destruction of nutrients, amino acids, and cell membranes in the microorganisms that make up biological soil crust. However, the biological soil crust has adapted to survive in very harsh environments with the aid of
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with algae in lichens. Free-living microfungi often function as decomposers, and contribute to soil microbial biomass. Many microfungi in biological soil crusts have adapted to the intense light conditions by evolving the ability to produce
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Soils in arid regions are slow-forming and easily eroded. Crust organisms contribute to increased soil stability where they occur. Cyanobacteria have filamentous growth forms that bind soil particles together, and hyphae of fungi and
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of the water. Soils that were less than 80% sand had greater infiltration due to biocrust creating soil aggregates. Other factors like plant roots may play a role in water retention and soil moisture at depths below the soil crust.
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in soil crusts are present just below the soil surface where they are partially protected from UV radiation. They become inactive when dry and reactivate when moistened. They can photosynthesize to fix carbon from the atmosphere.
1502:; Leslie, Alexander D.; Rodriguez-Caballero, Emilio; Zhang, Yuanming; Barger, Nichole N. (November 2019). Vries, Franciska (ed.). "Towards a predictive framework for biocrust mediation of plant performance: A meta-analysis".
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water content was greater than 60% whereas green algae showed little response to high water content. Photosynthesis rates are also dependent on temperature, with rates increasing up to approximately 28 °C (82 °F).
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radiation present in areas where biological soil crusts occur, biological soil crusts appear darker than the crustless soil in the same area due to the UV-protective pigmentation of cyanobacteria and other crust organisms.
452:, the abundance of lichens and mosses in biological soil crusts generally increases with increasing clay and silt content and decreasing sand. Also, habitats that are more moist generally support more lichens and mosses.
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The biological soil crust is an integral part of many arid and semi-arid ecosystems as an essential contributor to conditions such as dust control, water acquisition, and contributors of soil nutrients. Biocrust is
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was found to exist in a dormant, metabolically inactive state beneath the surface of the crust in periods of drought or water deficiency. When the biocrust eventually receives precipitation, it is able to perform
241:. These filaments bind soil particles throughout the uppermost soil layers, forming a 3-D net-like structure that holds the soil together in a crust. Other common cyanobacteria species are as those in the genus
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Angel, Roey; Conrad, Ralf (June 2013). "Elucidating the microbial resuscitation cascade in biological soil crusts following a simulated rain event: Microbial resuscitation in biological soil crusts".
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for plants that grow near biological soil crusts. This can occur through N fixation by cyanobacteria in the crusts, increased trapment of nutrient-rich dust, as well as increased concentrations of
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Deines, Lynell; Rosentreter, Roger; Eldridge, David J.; Serpe, Marcelo D. (June 2007). "Germination and seedling establishment of two annual grasses on lichen-dominated biological soil crusts".
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Estimates for annual carbon inputs range from 0.4 to 37 g/cm*year depending on successional state. Estimates of total net carbon uptake by crusts globally are ~3.9 Pg/year (2.1–7.4 Pg/year).
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Naylor, Dan; Sadler, Natalie; Bhattacharjee, Arunima; Graham, Emily B.; Anderton, Christopher R.; McClure, Ryan; Lipton, Mary; Hofmockel, Kirsten S.; Jansson, Janet K. (17 October 2020).
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in varying proportions. These organisms live in intimate association in the uppermost few millimeters of the soil surface, and are the biological basis for the formation of soil crusts.
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with more "leafy" structures that can be attached to the soil at only one portion. Lichens with algal symbionts can fix atmospheric carbon, while lichens with cyanobacterial symbionts
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Cyanobacteria are primarily responsible for the pigment and rejuvenation of the crust during environmental changes that result in short spurts of rehydration for the biocrust.
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Biological soil crusts do not compete with vascular plants for nutrients, but rather have been shown to increase nutrient levels in plant tissues, which results in higher
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Green, T. G. Allan; Proctor, Michael C. F. (2016). "Physiology of
Photosynthetic Organisms within Biological Soil Crusts: Their Adaptation, Flexibility, and Plasticity".
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Kheirfam, Hossein; Sadeghi, Seyed
Hamidreza; Homaee, Mehdi; Zarei Darki, Behrouz (2017). "Quality improvement of an erosion-prone soil through microbial enrichment".
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of lichens and mosses also have similar effects. The increased surface roughness of crusted areas compared to bare soil further improves resistance to wind and water
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Rosentreter, R., M. Bowker, and J. Belnap. 2007. A Field Guide to
Biological Soil Crusts of Western U.S. Drylands. U.S. Government Printing Office, Denver, Colorado.
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0.7–100 kg/ha per year, from hot deserts in
Australia to cold deserts. Estimates of total biological nitrogen fixation are ~ 49 Tg/year (27–99 Tg/year).
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Havrilla, Caroline A.; Chaudhary, V. Bala; Ferrenberg, Scott; Antoninka, Anita J.; Belnap, Jayne; Bowker, Matthew A.; Eldridge, David J.; Faist, Akasha M.;
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Cantón, Yolanda; Chamizo, Sonia; Rodriguez-Caballero, Emilio; Lázaro, Roberto; Roncero-Ramos, Beatriz; Román, José Raúl; Solé-Benet, Albert (6 March 2020).
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cyanobacteria, supposed as a new step, biologic, sustainable, eco-friendly and economically-effective technique to rehabilitate biological soil crust.
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including coarse litter application (such as straw) and planting vascular plants, but these are costly and labor-intensive techniques. Spraying
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cannot compete for light with vascular plants. Across the globe, biological soil crusts can be found on all continents including
Antarctica.
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years) in coarse soil textured, dry environments. Recovery times also depend on disturbance regime, site, and availability of propagules.
1028:"Carbon and nitrogen fixation differ between successional stages of biological soil crusts in the Colorado Plateau and Chihuahuan Desert"
547:. Aggregates of soil formed by crust organisms also increase soil aeration and provide surfaces where nutrient transformation can occur.
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and other disturbances and can require long time periods to recover composition and function. Biological soil crusts are also known as
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A recent study in China shows that biocrusts have been an import factor in the preservation of sections of the Great Wall built using
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awakened from dormancy. Cyanobacteria are able to repeat this process over and over again in the event of rehydration in the future.
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Bowker, Matthew A. (March 2007). "Biological Soil Crust
Rehabilitation in Theory and Practice: An Underexploited Opportunity".
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Rammed-earth section of the Great Wall of China. Research shows that biocrust is a natural factor in preserving the structure.
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populations also increase with more developed crusts due to increased microhabitats produced by the crust microtopography.
1830:"Controlling rainfall-induced soil loss from small experimental plots through inoculation of bacteria and cyanobacteria"
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Abed, Raeid M. M.; Polerecky, Lubos; Al-Habsi, Amal; Oetjen, Janina; Strous, Marc; de Beer, Dirk (6 November 2014).
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Belnap, Jayne (15 October 2006). "The potential roles of biological soil crusts in dryland hydrologic cycles".
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The increased nutrient status of plant tissue in areas where biological soil crusts occur can directly benefit
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The presence of biological soil crust cover can differentially inhibit or facilitate plant seed catchment and
1310:"Rapid Recovery of Cyanobacterial Pigments in Desiccated Biological Soil Crusts following Addition of Water"
1457:"Water Regulation in Cyanobacterial Biocrusts from Drylands: Negative Impacts of Anthropogenic Disturbance"
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of these surfaces cause microtopography such as rolling hills and steep pinnacles. Due to the intense
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crusts occur in areas where higher precipitation results in increased cover of lichen and mosses, and
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mosses. Liverworts can be flat and ribbon-like or leafy. They can reproduce by spore formation or by
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1646:"Climate change and physical disturbance cause similar community shifts in biological soil crusts"
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Elbert, W.; Weber, B.; Burrows, S.; Steinkamp, J.; Budel, B.; Andreae, M. O.; Poschl, U. (2012).
1252:"Dynamic cyanobacterial response to hydration and dehydration in a desert biological soil crust"
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The increased surface roughness associated with biological soil crusts increase the capture of
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Kheirfam, Hossein; Sadeghi, Seyed
Hamidreza; Zarei Darki, Behrouz; Homaee, Mehdi (May 2017).
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Biological Soil Crusts. (2014, July 17). Garcia-Pichel Lab. Retrieved March 20, 2023, from
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gel has been tried but this has adversely affected photosynthesis and nitrogen fixation of
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Solhaug, Knut Asbjørn; Gauslaa, Yngvar; Nybakken, Line; Bilger, Wolfgang (April 2003).
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varies by crust composition because only cyanobacteria and cyanolichens fix nitrogen.
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237:. These species form bundled filaments that are surrounded by a gelatinous sheath of
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1705:"Response of desert biological soil crusts to alteration in precipitation frequency"
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Evans, R. D.; Johansen, J. R. (1999). "Microbiotic Crusts and
Ecosystem Processes".
247:, which can also form sheaths and sheets of filaments that stabilize the soil. Some
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Communities of living organisms on the soil surface in arid and semi-arid ecosystems
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Belnap, Jayne (May 2003). "The world at your feet: desert biological soil crusts".
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lichens with scale- or plate-like bodies that are raised above the soils, and
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to the negatively charged clay particles bound by cyanobacterial filaments.
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983:"Soil Microbiomes Under Climate Change and Implications for Carbon Cycling"
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1135:"The arid environments: Arid zone soils and importance of soil properties"
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affects biological soil crusts by altering the timing and magnitude of
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135:. Biological soil crusts perform important ecological roles including
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Ferrenberg, Scott; Reed, Sasha C.; Belnap, Jayne (September 2015).
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in biological soil crusts can occur as free-living species, or in
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Bowker, Matthew A., Manuel
Delgado-Baquerizo (8 December 2023).
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Housman, D.C.; Powers, H.H.; Collins, A.D.; Belnap, J. (2006).
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The darkened surfaces of biological soil crusts decreases soil
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https://garcia-pichel.lab.asu.edu/labo/biological-soil-crusts/
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Lichens are often distinguished by growth form and by their
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Biological Soil Crusts: An
Organizing Principle in Drylands
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10.1890/1540-9295(2003)001[0181:TWAYFD]2.0.CO;2
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species are also able to fix atmospheric nitrogen gas into
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Biological soil crusts are formed in open spaces between
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1584:"Biocrusts protect the Great Wall of China from erosion"
1211:. Ecological Studies. Vol. 226. pp. 347–381.
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Food and Agriculture Organization of the United Nations
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159:. They can be damaged by fire, recreational activity,
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897:"UV-induction of sun-screening pigments in lichens"
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111:. They are found throughout the world with varying
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196:Biological soil crusts are most often composed of
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1140:Arid zone forestry: A guide for field technicians
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564:and flat, infiltration rates can be decreased by
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939:"Biological Soil Crusts: Ecology and Management"
872:. Southern Arizona Desert Botany. Archived from
839:"Cryptobiotic Soils: Holding the Place in Place"
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1650:Proceedings of the National Academy of Sciences
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600:, and thus increase both the fertility and the
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643:and appears to resurrect. In this stage, the
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1703:Belnap, J; Phillips, SL; Miller, ME (2004).
1629:: CS1 maint: multiple names: authors list (
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956:. Technical Reference 1730-2. Archived from
661:Biological soil crust role in soil hydrology
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525:Geophysical and geomorphological properties
512:Biological soil crust contributions to the
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96:are communities of living organisms on the
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987:Annual Review of Environment and Resources
870:"Cryptobiotic Crust in the Sonoran Desert"
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488:Biological soil crusts contribute to the
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281:. Mosses are usually classified as short
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555:The effect of biological soil crusts on
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1000:10.1146/annurev-environ-012320-082720
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634:A filamentous cyanobacterium called
27:Cryptobiotic soil, cryptogamic soil,
1886:, USGS Canyonlands Research Station
937:Belnap, Jayne; et al. (2001).
868:Moore, Lorena B. (March 23, 2010).
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29:microbiotic soil, microphytic soil,
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1109:Critical Reviews in Plant Sciences
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371:can bind soil particles together.
273:Bryophytes in soil crusts include
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424:Natural Bridges National Monument
1871:
1772:10.1111/j.1526-100X.2006.00185.x
914:10.1046/j.1469-8137.2003.00708.x
837:Belnap, Jayne (August 5, 2013).
674:Role in the biological community
608:Hydration and dehydration cycles
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363:, and are called black fungi or
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946:U.S. Department of the Interior
474:Ecosystem function and services
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151:and water relations and affect
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1052:10.1016/j.jaridenv.2005.11.014
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339:as well. Lichens produce many
1:
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802:Removal of stressors such as
684:Germination and establishment
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1855:10.1016/j.catena.2017.01.006
1335:10.1371/journal.pone.0112372
1217:10.1007/978-3-319-30214-0_18
1032:Journal of Arid Environments
742:Human impacts and management
7:
1815:10.1016/j.still.2016.08.021
798:Conservation and management
44:Hovenweep National Monument
10:
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1374:Environmental Microbiology
734:species in the community.
679:Effects on vascular plants
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439:Variation throughout range
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1795:Soil and Tillage Research
1729:10.1007/s00442-003-1438-6
1561:10.1007/s11104-007-9256-y
1500:Huber-Sannwald, Elisabeth
1121:10.1080/07352689991309199
950:Bureau of Land Management
598:plant-essential nutrients
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422:Biological soil crust in
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42:Biological soil crust in
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1034:(Submitted manuscript).
393:Formation and succession
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311:. Crust lichens include
119:, soil characteristics,
1671:10.1073/pnas.1509150112
1516:10.1111/1365-2745.13269
1386:10.1111/1462-2920.12140
375:Free-living green algae
192:Biology and composition
155:and nutrient levels in
115:and cover depending on
1884:Biological soil crusts
1600:10.1126/sciadv.adk5892
1414:Hydrological Processes
954:U.S. Geological Survey
843:U.S. Geological Survey
756:
694:invasive plant species
602:water holding capacity
479:Biogeochemical cycling
431:
94:Biological soil crusts
1268:10.1038/ismej.2013.83
754:
636:Microcoleus vaginatus
628:Microcoleus vaginatus
421:
291:asexual fragmentation
22:Biological soil crust
1880:at Wikimedia Commons
696:such as cheatgrass (
551:Soil water relations
1846:2017Caten.152...40K
1807:2017STilR.165..230K
1760:Restoration Ecology
1721:2004Oecol.141..306B
1662:2015PNAS..11212116F
1656:(39): 12116–12121.
1426:2006HyPr...20.3159B
1326:2014PLoSO...9k2372A
1086:2012NatGe...5..459E
1044:2006JArEn..66..620H
445:species composition
133:disturbance regimes
113:species composition
1504:Journal of Ecology
757:
726:Effects on animals
557:water infiltration
432:
414:Geographical range
147:; they alter soil
145:soil stabilization
1878:Cryptobiotic soil
1876:Media related to
1474:10.3390/w12030720
1420:(15): 3159–3178.
1380:(10): 2799–3515.
1262:(11): 2178–2191.
1226:978-3-319-30212-6
1154:978-92-5-102809-4
1074:Nature Geoscience
774:off-road vehicles
767:Human disturbance
718:that are able to
518:Nitrogen fixation
319:lichens that are
141:nitrogen fixation
91:
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1588:Science Advances
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1510:(6): 2789–2807.
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1434:10.1002/hyp.6325
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508:Nitrogen cycling
337:can fix nitrogen
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849:on June 3, 2016
835:
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706:Nutrient levels
699:Bromus tectorum
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399:vascular plants
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349:
333:foliose lichens
305:
299:
285:mosses or tall
271:
265:
239:polysaccharides
228:
222:
194:
189:
187:Natural history
157:vascular plants
137:carbon fixation
125:plant community
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1866:External links
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809:polyacrylamide
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784:Climate change
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747:Human benefits
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615:poikilohydric
605:
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584:Dust-trapping
581:
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561:soil moisture
558:
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253:bio-available
250:
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226:Cyanobacteria
220:Cyanobacteria
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206:cyanobacteria
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1906:Soil biology
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1788:
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1763:
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1639:
1625:cite journal
1613:. Retrieved
1609:10261/340361
1591:
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1552:
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1525:10261/202886
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958:the original
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874:the original
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847:the original
812:
801:
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778:
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761:rammed earth
758:
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650:M. vaginatus
649:
645:M. vaginatus
644:
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503:
490:carbon cycle
487:
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450:climate zone
442:
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409:Distribution
403:
396:
384:
365:black yeasts
350:
323:to the soil
306:
272:
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242:
232:
229:
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181:cryptobiotic
180:
176:
172:
168:
164:
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92:
1801:: 230–238.
792:temperature
790:events and
690:germination
566:bioclogging
494:respiration
386:Green algae
381:Green algae
234:Microcoleus
177:microphytic
173:microbiotic
169:cryptogamic
153:germination
100:surface in
1900:Categories
1615:9 December
1467:(3): 720.
967:2014-03-24
907:: 91–100.
822:References
641:hydrotaxis
604:of soils.
352:Microfungi
329:squamulose
279:liverworts
263:Bryophytes
210:bryophytes
117:topography
109:ecosystems
82:bryophytes
1840:: 40–46.
1709:Oecologia
1534:202017609
1442:129013389
763:methods.
732:herbivore
367:. Fungal
356:symbiosis
325:substrate
321:appressed
287:perennial
269:Bryophyte
165:biocrusts
106:semi-arid
60:semi-arid
1916:Mycology
1780:51779646
1745:27306844
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1690:26371310
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1354:25375172
1314:PLOS ONE
1286:23739051
1147:. 1989.
592:. These
541:rhizoids
537:rhizines
492:through
341:pigments
317:areolate
313:crustose
31:biocrust
1911:Lichens
1842:Bibcode
1803:Bibcode
1717:Bibcode
1681:4593113
1658:Bibcode
1422:Bibcode
1345:4223047
1322:Bibcode
1277:3806265
1082:Bibcode
1040:Bibcode
880:May 10,
853:May 10,
814:Collema
804:grazing
720:chelate
712:biomass
594:Aeolian
545:erosion
469:Ecology
361:melanin
297:Lichens
257:ammonia
202:lichens
183:soils.
161:grazing
121:climate
74:lichens
66:Primary
52:Climate
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249:Nostoc
244:Nostoc
212:, and
167:or as
149:albedo
131:, and
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1530:S2CID
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1438:S2CID
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426:near
347:Fungi
214:algae
198:fungi
179:, or
86:algae
70:fungi
1733:PMID
1686:PMID
1631:link
1617:2023
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1350:PMID
1282:PMID
1221:ISBN
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952:and
882:2016
855:2016
590:dust
559:and
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315:and
277:and
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