Cyanobacteria form extensive macroscopic mats in shallow freshwater environments in the High Arctic and Antarctic. In these habitats, the communities are exposed to seasonal freezing and desiccation as well as to freeze-thawing and drying-rewetting cycles. Here, we characterized the annual cycles of two Phormidium communities in very shallow seepages located in central Svalbard. We observed the structure of the communities and the morphology, ultrastructure, metabolic activity, and viability of filaments and single cells. The communities overwintered as frozen mats, which were formed by long filaments enclosed in thick multilayered polysaccharide sheaths. No morphologically and/or ultrastructurally distinct spore-like cells were produced for surviving the winter, and the winter survival of the communities was not provided by a few resistant cells, which did not undergo visible morphological and ultrastructural transformations. Instead, a high proportion of cells in samples (85 %) remained viable after prolonged freezing. The sheaths were the only morphological adaption, which seemed to protect the trichomes from damage due to freezing and freeze-associated dehydration. The cells in the overwintering communities were not dormant, as all viable cells rapidly resumed respiration after thawing, and their nucleoids were not condensed. During the whole vegetative season, defined by the presence of water in a liquid state, the communities were constantly metabolically active and contained
