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The last decades of research led to a change in understanding of lichens that are now seen as self-sustaining micro-ecosystems, harboring diverse microbial organisms in tight but yet not fully understood relationships. Among the diverse interdependencies, the relationship between the myco- and photobiont is the most crucial, determining the shape, and ecophysiological properties of the symbiotic consortium. Roughly 10% of lichens associate with cyanobacteria as their primary photobiont, termed cyanolichens. Up to now, the diversity of cyanobionts of bipartite lichens resolved by modern phylogenetic approaches is restricted to the filamentous and heterocytous genera of the order Nostocales. Unicellular photobionts were placed in the orders Chroococcales, Pleurocapsales, and Chroococcidiopsidales. However, especially the phylogeny and taxonomy of the Chroococcidiopsidales genera remained rather unclear. Here we present new data on the identity and phylogeny of photobionts from cyanolichens of the genera  Gonohymenia ,  Lichinella ,  Peccania , and  Peltula  from a broad geographical range. A polyphasic approach was used, combining morphological and cultivation-depending characteristics (microscopy, staining techniques, life cycle observation, baeocyte motility, and nitrogen fixation test) with phylogenetic analyses of the 16S rRNA and 16S–23S ITS gene region. We found an unexpectedly high cyanobiont diversity in the cyanobacterial lichens of the order Lichinales, including two new genera and seven new species, all of which were not previously perceived as lichen symbionts. As a result, we describe the novel unicellular Chroococcidiopsidales genera  Pseudocyanosarcina  gen. nov. with the species  Pseudocyanosarcina phycocyania  sp. nov. (from  Peltula clavata , Australia) and  Compactococcus  gen. nov. with the species  Compactococcus sarcinoides  sp. nov. (from  Gonohymenia  sp., Australia) and the new Chroococcidiopsidales species  Aliterella compacta  sp. nov. (from  Peltula clavata , Australia),  Aliterella gigantea  sp. nov. (from  Peltula capensis ; South Africa),  Sinocapsa ellipsoidea  sp. nov. (from  Peccania cerebriformis , Austria), as well as the two new Nostocales species  Komarekiella gloeocapsoidea  sp. nov. (from  Gonohymenia  sp., Czechia) and  Komarekiella globosa  sp. nov. (from  Lichinella cribellifera , Canary Islands, Spain). Our study highlights the role of cyanolichens acting as a key in untangling cyanobacterial taxonomy and diversity. With this study, we hope to stimulate further research on photobionts, especially of rare cyanolichens.

Opening the Gap: Rare Lichens With Rare Cyanobionts – Unexpected Cyanobiont Diversity in Cyanobacterial Lichens of the Order Lichinales