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Many terrestrial surfaces are covered by photoautotrophic communities, which are capable of synthesizing their own food from inorganic substances using sunlight. According to an analysis of previously published data, these communities account for nearly half of the biological nitrogen fixation on land. Many terrestrial surfaces, including soils, rocks and plants, are covered by photoautotrophic communities, capable of synthesizing their own food from inorganic substances using sunlight as an energy source 1 , 2 . These communities, known as cryptogamic covers, comprise variable proportions of cyanobacteria, algae, fungi, lichens and bryophytes, and are able to fix carbon dioxide and nitrogen from the atmosphere 3 . However, their influence on global and regional biogeochemical cycling of carbon and nitrogen has not yet been assessed. Here, we analyse previously published data on the spatial coverage of cryptogamic communities, and the associated fluxes of carbon and nitrogen, in different types of ecosystem across the globe. We estimate that globally, cryptogamic covers take up around 3.9 Pg carbon per year, corresponding to around 7% of net primary production by terrestrial vegetation. We derive a nitrogen uptake by cryptogamic covers of around 49 Tg per year, suggesting that cryptogamic covers account for nearly half of the biological nitrogen fixation on land. We suggest that nitrogen fixation by cryptogamic covers may be crucial for carbon sequestration by plants.

Cyanobacteria associated with biological soil crusts (BSCs) have important attributes, such as nitrogen fixation and soil stabilisation. However, research on these organisms has been minimal, and their diversity and distribution throughout temperate Europe is currently unknown. The SCIN (Soil Crust International) project is a multidisciplinary research initiative that aims to achieve improved understanding of the BSCs of Europe, one facet being an investigation into the cyanobacterial communities of BSCs across a latitudinal gradient. Cyanobacteria assemblages were analysed by both morphological and molecular analysis. Two treatments were applied prior to DNA extraction, continued sample wetting and a dry sample process, and 16S ribosomal RNA (rRNA) amplicons were processed by Illumina MiSeq sequencing. The results reveal high and variable cyanobacterial diversity with each site showing a unique assemblage. Many common cyanobacterial genera, for example Nostoc and Microcoleus, were found in all sites but the abundances of different genera varied considerably. The polyphasic approach was found to be essential in recording the presence of important cyanobacteria that a single method itself did not highlight. The wet and dry treatments showed some differences in diversity, but mainly in abundance, this may suggest how cyanobacterial composition of BSCs changes with seasonal variability. Investigating the diversity and community composition of cyanobacteria associated with biological soil crusts in climatically different regions of Europe.

Habitat fragmentation and forest management have been considered to drastically alter the nature of forest ecosystems globally. However, much uncertainty remains regarding the causative mechanisms mediating temperate forest responses, such as forest physical environment and the structure of woody plant assemblages, regardless of the role these forests play for global sustainability. In this paper, we examine how both habitat fragmentation and timber exploitation via silvicultural operations affect these two factors at local and habitat spatial scales in a hyper-fragmented landscape of mixed beech forests spanning more than 1500 km2 in SW Germany. Variables were recorded across 57 1000 m2 plots covering four habitats: small forest fragments, forest edges within large control forests, as well as managed and unmanaged forest interior sites. As expected, forest habitats differed in disturbance level, physical conditions and community structure at plot and habitat scale. Briefly, diversity of plant assemblages differed across all forest habitats (highest in edge forests) and correlated with integrative indices of edge, fragmentation and management effects. Surprisingly, managed and unmanaged forests did not differ in terms of species richness at local spatial scale, but managed forests exhibited a clear signal of physical/floristic homogenization as species promoted by silviculture proliferated; i.e. impoverished communities at landscape scale. Moreover, functional composition of plant communities responded to the microclimatic regime within forest fragments, resulting in a higher prevalence of species adapted to these microclimatic conditions. Our results underscore the notion that forest fragmentation and silvicultural management (1) promote changes in microclimatic regimes, (2) alter the balance between light-demanding and shade-adapted species, (3) support diverse floras across forest edges, and (4) alter patterns of beta diversity. Hence, in human-modified landscapes edge-affected habitats can be recognized as biodiversity reservoirs in contrast to impoverished managed interior forests. Furthermore, our results ratify the role of unmanaged forests as a source of environmental variability, species turnover, and distinct woody plant communities.

Biological soil crusts (BSCs) are key components of polar ecosystems. These complex communities are important for terrestrial polar habitats as they include major primary producers that fix nitrogen, prevent soil erosion and can be regarded as indicators for climate change. To study the genus richness of microalgae and Cyanobacteria in BSCs, two different methodologies were employed and the outcomes were compared: morphological identification using light microscopy and the annotation of ribosomal sequences taken from metatranscriptomes. The analyzed samples were collected from Ny-Ålesund, Svalbard, Norway, and the Juan Carlos I Antarctic Base, Livingston Island, Antarctica. This study focused on the following taxonomic groups: Klebsormidiophyceae, Chlorophyceae, Trebouxiophyceae, Xanthophyceae and Cyanobacteria. In total, combining both approaches, 143 and 103 genera were identified in the Arctic and Antarctic samples, respectively. Furthermore, both techniques concordantly determined 15 taxa in the Arctic and 7 taxa in the Antarctic BSC. In general, the molecular analysis indicated a higher microalgal and cyanobacterial genus richness (about 11 times higher) than the morphological approach. In terms of eukaryotic algae, the two sampling sites displayed comparable genus counts while the cyanobacterial genus richness was much higher in the BSC from Ny-Ålesund. For the first time, the presence of the genera Chloroidium, Ankistrodesmus and Dunaliella in polar regions was determined by the metatranscriptomic analysis. Overall, these findings illustrate that only the combination of morphological and molecular techniques, in contrast to one single approach, reveals higher genus richness for complex communities such as polar BSCs.

Biocrusts are associations of various prokaryotic and eukaryotic microorganisms in the top millimeters of soil, which can be found in every climate zone on Earth. They stabilize soils and introduce carbon and nitrogen into this compartment. The worldwide occurrence of biocrusts was proven by numerous studies in Europe, Africa, Asia and North America, leaving South America understudied. Using an integrative approach, which combines morphological and molecular characters (small subunit rRNA and ITS region), we examined the diversity of key biocrust photosynthetic organisms at four sites along the latitudinal climate gradient in Chile. The most northern study site was located in the Atacama Desert (arid climate), followed by open shrubland (semiarid climate), a dry forest region (Mediterranean climate) and a mixed broad leaved-coniferous forest (temperate climate) in the south. The lowest species richness was recorded in the desert (18 species), whereas the highest species richness was observed in the Mediterranean zone (40 species). Desert biocrusts were composed exclusively of single-celled Chlorophyta algae, followed by cyanobacteria. Chlorophyta, Streptophyta and cyanobacteria dominated semiarid biocrusts, whereas Mediterranean and temperate Chilean biocrusts were composed mostly of Chlorophyta, Streptophyta and Ochrophyta. Our investigation of Chilean biocrust suggests high biodiversity of South American biocrust phototrophs.

Biological soil crusts (BSCs) occur in arid and semi-arid regions worldwide including the Polar Regions. They are important ecosystem engineers, and their composition and areal coverage should be understood before assessing key current functional questions such as their role in biogeochemical nutrient cycles and possible climate change scenarios. Our aim was to investigate the variability of BSCs from Arctic Svalbard and the Antarctic Island, Livingston, using vegetation surveys based on classification by functional group. An additional aim was to describe the structure of BSCs and represent a classification system that can be used in future studies to provide a fast and efficient way to define vegetation type and areal coverage. Firstly, this study demonstrates huge areas occupied by BSCs in Arctic Svalbard, with up to 90 % of soil surface covered, dominated by bryophytes and cyanobacteria, and showing an unexpectedly high variability in many areas. Livingston Island has lower percentage coverage, up to 55 %, but is dominated by lichens. Our findings show that both Polar Regions have varied BSC coverage, within the sites and between them, especially considering their harsh climates and latitudinal positions. Secondly, we have classified the BSCs of both areas into a system that describes the dominant functional groups and local geography, creating a simple scheme that allows easy identification of the prevailing vegetation type. Our results represent the first contribution to the description of BSCs based on their functional group composition in Polar Regions.

Plants and other organisms have evolved structures and mechanisms for colonizing land since the Early Ordovician. In this context, their surfaces, the crucial physical interface with the environment, are mainly considered barriers against water loss. It is suggested that extreme water repellency (superhydrophobicity) was an additional key innovation for the transition of algae from water to land some 400 mya. Superhydrophobicity enhances gas exchange on land and excludes aquatic competitors in water films. In a different context, in material science and surface technology, superhydrophobicity has also become one of the most important bioinspired innovations enabling the avoidance of water films and contamination. Here, we present data for an extremely water-repellent cyanobacterial biofilm of the desiccation tolerant Hassallia byssoidea providing evidence for a much earlier prokaryotic Precambrian (ca. 1–2 bya) origin of superhydrophobicity and chemical heterogeneities associated with land transition. The multicellular cyanobacterium is functionally differentiated in a submerged basal hydrophilic absorbing portion like a “rhizoid” and an upright emersed superhydrophobic “phyllocauloid” filament for assimilation, nitrogen fixation, and splash dispersed diaspores. Additional data are provided for superhydrophobic surfaces in terrestrial green algae and in virtually all ancestral land plants (Bryophytes, ferns and allies, Amborella , Nelumbo ), slime molds, and fungi. Rethinking of superhydrophobicity as an essential first step for life in terrestrial environments is suggested.

Unraveling the complex relationship between lichen fungal and algal partners has been crucial in understanding lichen dispersal capacity, evolutionary processes, and responses in the face of environmental change. However, lichen symbiosis remains enigmatic, including the ability of a single fungal partner to associate with various algal partners. Psora decipiens is a characteristic lichen of biological soil crusts (BSCs), across semi‐arid, temperate, and alpine biomes, which are particularly susceptible to habitat loss and climate change. The high levels of morphological variation found across the range of Psora decipiens may contribute to its ability to withstand environmental change. To investigate Psora decipiens acclimation potential, individuals were transplanted between four climatically distinct sites across a European latitudinal gradient for 2 years. The effect of treatment was investigated through a morphological examination using light and SEM microscopy; 26S rDNA and rbcL gene analysis assessed site‐specific relationships and lichen acclimation through photobiont switching. Initial analysis revealed that many samples had lost their algal layers. Although new growth was often determined, the algae were frequently found to have died without evidence of a new photobiont being incorporated into the thallus. Mycobiont analysis investigated diversity and determined that new growth was a part of the transplant, thus, revealing that four distinct fungal clades, closely linked to site, exist. Additionally, P. decipiens was found to associate with the green algal genus Myrmecia, with only two genetically distinct clades between the four sites. Our investigation has suggested that P. decipiens cannot acclimate to the substantial climatic variability across its environmental range. Additionally, the different geographical areas are home to genetically distinct and unique populations. The variation found within the genotypic and morpho‐physiological traits of P. decipiens appears to have a climatic determinant, but this is not always reflected by the algal partner. Although photobiont switching occurs on an evolutionary scale, there is little evidence to suggest an active environmentally induced response. These results suggest that this species, and therefore, other lichen species, and BSC ecosystems themselves may be significantly vulnerable to climate change and habitat loss. We have shown that Psora decipiens, a characteristic and important lichen of biological soil crusts worldwide, cannot acclimate to the substantial climatic variability across its environmental range. Different geographical areas are home to genetically distinct and unique populations. Although photobiont switching occurs on an evolutionary scale there is little evidence to suggest an active environmentally induced response. These results suggest that this species, and therefore, other lichen species and BSC ecosystems themselves, may be significantly vulnerable to climate change and habitat loss.

Biological soil crusts (BSCs) are found in all dryland regions of the world, including the polar regions. They are also known to occur in the southern African region. Although there were a number of case studies on BSCs from that region, we did not know if they are a normal part of the vegetation cover or just a phenomenon that occasionally occurs here and there. In order to investigate diversity, distribution patterns, and the driving factors of both, we followed a random sampling system of observatories along a transect, stretching from the Namibian-Angolan border down south to the Cape Peninsula, covering seven different major biomes. Biological soil crusts were found to occur in six out of seven biomes. Despite the fact that soil-dwelling algae occurred in the Fynbos biome, crust formation was not observed for hitherto unknown reasons. Seven BSC types were distinguished on the basis of morphology and taxonomic composition: three of them were cyanobacteria-dominated, one with additional chlorolichens, two with bryophytes, one hypolithic type restricted to quartz gravel pavements, and the unique lichen fields of the Namib Desert. Besides 29 green algal species in 21 genera, one heterokont alga, 12 cyanolichens, 14 chlorolichens, two genera of liverworts, and three genera of mosses, these crusts are positioned among the most diverse BSCs worldwide mainly because of the unusual high cyanobacterial species richness comprising 58 species in 21 genera. They contribute considerably to the biodiversity of arid and semi-arid bioregions. Taxonomic diversity of cyanobacteria was significantly higher in the winter rain zone than in the summer rain zone (54 versus 32 species). The soil photosynthetic biomass (chlorophylla/m²), the carbon content of the soil and the number of BSC types were significantly higher in the winter rain zone (U ₂₇, ₂₉ = 215.0, p = 0.004 [chla]; U ₂₁, ₂₁ = 135.0, p = 0.031 [C]; U ₂₇, ₂₉ = 261.5, p = 0.028 [BSC types]; excluding the fog-dominated Namib biome). The winter rain zone is characterized by a lower precipitation amount, but a higher rain frequency with the number of rainy days more evenly distributed over the year. The dry period is significantly shorter per year in the winter rain zone (U ₈, ₉ = 5.0, p = 0.003). We conclude that rain frequency and duration of dry periods rather than the precipitation amount is the main factor for BSC growth and succession. Nitrogen content of the soils along the transect was generally very low and correlated with soil carbon content. There was a weak trend that an increasing proportion of silt and clay (<0.63 mm) in the soil is associated with higher values of BSC chlorophyll content (Pearson correlation coefficient = 0.314, p = 0.237). Furthermore, we found a significant positive correlation between silt and clay and the number of BSC types (Pearson correlation coefficient = 0.519, p = 0.039), suggesting that fine grain-size promotes BSC succession and their biomass content. Lichens and bryophytes occurred in BSCs with lower disturbance frequencies (e.g. trampling) only. Crust thickness and chlorophyll content increased significantly from crusts of the early successional type to the late successional crust types. From our results, we conclude that BSCs are a normal and frequent element of the vegetation in arid and semi-arid southwestern Africa, and that rain frequency and duration of dry periods rather than the precipitation amount are the key factors for the development, differentiation and composition of BSCs.