Biological soil crust

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.

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 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 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 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.

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.