Controls on distribution patterns of biological soil crusts at micro- to global scales
Biocrusts are heterogeneously distributed in space. The drivers of their distribution patterns vary depending on the spatial scale of observation. Globally, there are about 1337 cyanobacteria, algae, bryophyte, and lichen species reported as components of biocrusts. At the broadest biogeographical scales, the degree and age of isolation of land masses may dictate distribution of these species and the similarities of the floras of different continents. At intra-continental and smaller scales, climate strongly influences abundance and community composition of biocrusts. Within drylands, biocrusts become more abundant as precipitation increases. The seasonality of rainfall is about equally important, with regions receiving most precipitation as winter rain and snow exhibiting the highest abundance and greatest relative cover of bryophyte and lichens vs. cyanobacteria. Temperature gradients may dictate the dominant cyanobacterium present in the community. At eco-regional and smaller scales, edaphic gradients determined by either soil parent materials or geomorphology or both become particularly influential. Globally, the most significant soil properties influencing the eco-regional scale cover and richness of biocrusts in dryland environments are soil texture, pH, and soil CaCO3 content. Sandier soils tend to favour development of cyanobacterial biocrusts, whereas mosses and lichens tend to be more abundant on finer textured soils. The alkalinity and CaCO3 content of soils are associated with greater bioocrust abundance in some regions, and dictates the species composition in the bryophyte and lichen component. Globally, gypsiferous soils are often associated with distinct floras and high abdundances of biocrusts, especially lichens. At local to micro-scales, biocrusts often are better developed in habitats with lower radiation loads such as polar-oriented slopes, or shaded habitats. Also at small scales, vascular plant canopies buffer microclimate for biocrusts, but also exert negative influences such as burial by litter. While our knowledge of biocrust distribution has advanced rapidly, there are considerable geographic and taxonomic gaps in our knowledge and a pronounced lack of truly global studies.