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Life at the atmosphere-lithosphere boundary is an ancient terrestrial niche that is sparsely covered by thin subaerial biofilms. The microbial inhabitants of these biofilms (a) have adapted to all types of terrestrial/subaerial stresses (e.g., desiccation, extreme temperatures, low nutrient availability, intense solar radiation), (b) interact with minerals that serve as both a dwelling and a source of mineral nutrients, and (c) provoke weathering of rocks and soil formation. Subaerial communities comprise heterotrophic and phototrophic microorganisms that support each other's lifestyle. Major lineages of eubacteria associated with the early colonization of land (e.g., Actinobacteria, Cyanobacteria) are present in these habitats along with eukaryotes such as microscopic green algae and ascomycetous fungi. The subaerial biofilm inhabitants have adapted to desiccation, solar radiation, and other environmental challenges by developing protective, melanized cell walls, assuming microcolonial architectures and symbiotic lifestyles. How these changes occurred, their significance in soil formation, and their potential as markers of climate change are discussed below.
The class Dothideomycetes (along with Eurotiomycetes) includes numerous rock-inhabiting fungi (RIF), a group of ascomycetes that tolerates surprisingly well harsh conditions prevailing on rock surfaces. Despite their convergent morphology and physiology, RIF are phylogenetically highly diverse in Dothideomycetes. However, the positions of main groups of RIF in this class remain unclear due to the lack of a strong phylogenetic framework. Moreover, connections between rock-dwelling habit and other lifestyles found in Dothideomycetes such as plant pathogens, saprobes and lichen-forming fungi are still unexplored. Based on multigene phylogenetic analyses, we report that RIF belong to Capnodiales (particularly to the family Teratosphaeriaceae s.l.), Dothideales, Pleosporales, and Myriangiales, as well as some uncharacterised groups with affinities to Dothideomycetes. Moreover, one lineage consisting exclusively of RIF proved to be closely related to Arthoniomycetes, the sister class of Dothideomycetes. The broad phylogenetic amplitude of RIF in Dothideomycetes suggests that total species richness in this class remains underestimated. Composition of some RIF-rich lineages suggests that rock surfaces are reservoirs for plant-associated fungi or saprobes, although other data also agree with rocks as a primary substrate for ancient fungal lineages. According to the current sampling, long distance dispersal seems to be common for RIF. Dothideomycetes lineages comprising lichens also include RIF, suggesting a possible link between rock-dwelling habit and lichenisation.
Solar energy has been transformed into useful redox differences or disequilibria within the Earth´s crust since the onset of anoxygenic and oxygenic photosynthesis in the Precambrian. Inorganic oxidized carbon is transformed into reduced carbon compounds by capturing and storing solar energy. During this process, many different organic compounds are formed including carbohydrates, proteins, hydrocarbons, and various other complex organic metabolic products and their diagenetic polymerization products (melanin, humic substances, petroleum, coal, and kerogen). Many of these solar energy-enriched compounds, however, are oxidized immediately or during the diagenetic transformation of sediments. The oxidation agents are oxygen, sulfate, iron, and other oxidized compounds, which in turn are partly enriched with the original solar energy. On a global biogeochemical scale, however, sulfur and iron are the most important elements. Geological evidence shows that biogeochemical cycles tend to yield stable ratios between the most oxidized forms of carbon (carbon dioxide and calcium/magnesium carbonate) and the most reduced forms (diamond, coal, methane, and hydrocarbons). Throughout the Earths history and evolution, this equilibrium ratio is around 1:4, maximally 1:5. When too much carbon is stored in the crust in the form of reduced compounds or vice versa, climatic and biogeomorphogenetic consequences upset the equilibrium. The biosphere reacts in a way to return to the optimal ratio. Excellent examples for this fluctuating equilibrium are the Carboniferous (too much organic carbon stored), the Permian (too little organic carbon stored), and the Tertiary with a generally equivalent production of hydrocarbons and carbonates. At present, we are in a period in which there is a global biogeochemical need to oxidize reduced carbon compounds as fast as possible in order to avoid even more dramatic global climate shifts. The highly evolved human genome seems to be the tool for this shift. Enormous amounts of reduced carbon are turned into the oxidized form as carbon dioxide, which by various biogeochemical pathways is quickly transformed into carbonate, another oxidized form of carbon that can be stabilized and stored in the sedimentary record. Fast recycling of excessively stored solar energy may enable the survival of a global biosphere under highly stressed conditions.