Climatic factors, soil chemistry and geography are considered as major factors affecting lichen distribution and diversity. To determine how these factors limit or support the associations between the symbiotic partners, we revise the lichen symbiosis as a network of relationships here. More than one thousand thalli of terricolous Cladonia lichens were collected at sites with a wide range of soil chemical properties from seven biogeographical regions of Europe. A total of 18 OTUs of the algal genus Asterochloris and 181 OTUs of Cladonia mycobiont were identified. We displayed all realized pairwise mycobiont–photobiont relationships and performed modularity analysis. It revealed four virtually separated modules of cooperating OTUs. The modules differed in mean annual temperature, isothermality, precipitation, evapotranspiration, soil pH, nitrogen, and carbon contents. Photobiont switching was strictly limited to algae from one module, i.e., algae of similar ecological preferences, and only few mycobionts were able to cooperate with photobionts from different modules. Thus, Cladonia mycobionts generally cannot widen their ecological niches through photobiont switching. The modules also differed in the functional traits of the mycobionts, e.g., sexual reproduction rate, presence of soredia, and thallus type. These traits may represent adaptations to the environmental conditions that drive the differentiation of the modules. In conclusion, the promiscuity in Cladonia mycobionts is strictly limited by climatic factors and soil chemistry.

Lichenised algae and cyanobacteria are known to be shared and selected by unrelated lichen-forming fungi coexisting in so called photobiont-mediated guilds. Life in such a guild could be crucial for the survival of a large group of lichen fungi dependent on horizontal transmission of photobionts. Here, we investigate frequent lichen phycobionts of the genus Trebouxia in rock-dwelling lichen communities. We found intensive and repeated sharing of specific Trebouxia assemblages by co-occurring lichens across distant localities. Rock chemistry, expressed as pH, determined the composition of photobiont pools and separate three saxicolous lichen guilds, sharing environmentally specific photobiont groups. Moreover, unlike the majority of lichen fungi, many Trebouxia photobionts represent opportunists in the choice of general substrate form (soil– rock–tree bark/wood), maintaining their pH preferences. Thus, saxicolous communities form just a part of a complex guild system which is in principle mediated by environmentally conditioned groups of naturally co-occurring photobionts. The complexity of the system is influenced by diverse photobiont life strategies, including also dispersal style. The findings of photobionts strictly or predominantly associated with sexually reproducing fungi stimulate us to emphasise the role of free-dispersing photobionts in the establishment and maintenance of lichen guilds.

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River gravel bars are dynamic and heterogeneous habitats standing on transition between aquatic and terrestrial environment. Periodical flooding, low nutrient content, frost, missing safe sites, drought, and heat on the ground surface significantly influence life in these habitats. Mutualistic symbiosis may be a successful strategy for organisms to survive and to proliferate under harsh conditions. The lichen genus Stereocaulon was selected as a model symbiotic system among the organisms living on river gravel bars. The aim of our work was to determine effect of this dynamic environment on a phycobiont (i.e., green eukaryotic photobiont) community structure. We analysed 147 Stereocaulon specimens collected in the Swiss Alps using Sanger sequencing (fungal ITS rDNA, algal ITS rDNA, algal actin type I gene) and 8 selected thalli and 12 soil samples using Illumina metabarcoding (ITS2 rDNA). We performed phytosociological sampling on each study plot (n=13). Our analyses of communities of phycobionts, lichens, bryophytes, and vascular plants indicated an ongoing colonisation by phycobionts and gradual change of phycobiont community along to the successional gradient. We recovered great phycobiont diversity associated with Stereocaulon mycobionts including algae reported as phycobionts for the first time. Each of two Stereocaulon mycobiont OTUs has distinct pool of predominant phycobionts in the study area. Finally, all thalli selected for Illumina metabarcoding contained a wide range of additional intrathalline algae, i.e., showed algal plurality. In general, succession process on newly emerged or recently disturbed localities also takes place within a community of microscopic symbiotic organisms, such as phycobionts.

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