Communities of living organisms on the soil surface in arid and semi-arid ecosystems
Biological soil crusts, often abbreviated as biocrusts, are communities of living organisms inhabiting the surface of soils in arid and semi-arid ecosystems, which form stable aggregates of soil particles in a thin layer millimeters to centimeters thick. They are found throughout the world with varying species composition and cover depending on topography, soil characteristics, climate, plant community, microhabitats, and disturbance regimes. An estimated 12% of Earth's surface is covered by biocrusts. 
Biological soil crusts perform important ecological roles including carbon fixation, nitrogen fixation and soil stabilization; they alter soil albedo and water relations and affect germination and nutrient levels in vascular plants. They can be damaged by fire, recreational activity, grazing and other disturbances and can require long time periods to recover composition and function. Other names for biological soil crusts include "cryptogamic", "microbiotic", "microphytic", or "cryptobiotic" soils.
Natural history.
Biology and composition.
Biological soil crusts are most often composed of fungi, lichens, cyanobacteria, bryophytes, and algae 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.
Cyanobacteria.
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 "Microcoleus". These species form bundled filaments that are surrounded by a gelatinous sheath of polysaccharides. 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 "Nostoc", which can also form sheaths and sheets of filaments that stabilize the soil. Some "Nostoc" species are also able to fix atmospheric nitrogen gas into bio-available forms such as ammonia.
Bryophytes.
Bryophytes in soil crusts include mosses and liverworts. Mosses are usually classified as short annual mosses or tall perennial mosses. Liverworts can be flat and ribbon-like or leafy. They can reproduce by spore formation or by asexual fragmentation, and photosynthesize to fix carbon from the atmosphere.
About 250 moss species have been recorded in biocrust communities, mostly from the families Bryaceae, Pottiaceae, and Grimmiaceae.
Lichens.
Lichens are often distinguished by growth form and by their photosymbiont. Crust lichens include crustose and areolate lichens that are appressed to the soil substrate, squamulose lichens with scale- or plate-like bodies that are raised above the soils, and foliose lichens 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 can fix nitrogen as well. Lichens produce many pigments that help protect them from radiation.
Fungi.
Microfungi in biological soil crusts can occur as free-living species, or in symbiosis 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 melanin, and are called black fungi or black yeasts. Fungal hyphae can bind soil particles together.
Free-living green algae.
Green algae 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.
Formation and succession.
Biological soil crusts are formed in open spaces between vascular plants. 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.
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 years) in coarse soil textured, dry environments. Recovery times also depend on disturbance regime, site, and availability of propagules.
Distribution.
Geographical range.
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 cannot compete for light with vascular plants. Across the globe, biological soil crusts can be found on all continents including Antarctica.
Variation throughout range.
The species composition 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 climate zone, 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.
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 crusts occur in areas where higher precipitation results in increased cover of lichen and mosses, and frost heaving of these surfaces cause microtopography such as rolling hills and steep pinnacles. Due to the intense UV 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.
Ecology.
Ecosystem function and services.
Biogeochemical cycling.
Carbon cycling.
Biological soil crusts contribute to the carbon cycle through respiration and photosynthesis 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 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 .
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).
Nitrogen cycling.
Biological soil crust contributions to the nitrogen cycle varies by crust composition because only cyanobacteria and cyanolichens fix nitrogen. Nitrogen fixation 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 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).
Geophysical and geomorphological properties.
Soil stability.
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 rhizines/rhizoids 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 erosion. Aggregates of soil formed by crust organisms also increase soil aeration and provide surfaces where nutrient transformation can occur.
Soil water relations.
The effect of biological soil crusts on water infiltration and soil moisture 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 and flat, infiltration rates can be decreased by bioclogging.
Albedo.
The darkened surfaces of biological soil crusts decreases soil albedo (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.
Dust-trapping.
The increased surface roughness associated with biological soil crusts increase the capture of dust. These Aeolian deposits of dust are often enriched in plant-essential nutrients, and thus increase both the fertility and the water holding capacity of soils.
Hydration and dehydration cycles.
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 poikilohydric 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, 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.
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 cyanobacteria. 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 "Microcoleus vaginatus" 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. 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.
A filamentous cyanobacterium called "Microcoleus vaginatus" 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 hydrotaxis and appears to resurrect. In this stage, the "M. vaginatus" 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 "M. vaginatus" 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 awakened from dormancy. Cyanobacteria are able to repeat this process over and over again in the event of rehydration in the future.
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.
Biological soil crust role in soil hydrology.
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 the absorption and retention of moisture in the soil. In arid and semi-arid environments biocrust can cover