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lichen eukaryotes bark peltigeraceae mycobiont moss orzega ascomycete fungi phycobiont photobiont spruce ecology lichenes lichens mushrooms lecanoromycetes belonging karelia bryophyta mosses lobariaceae forest thallus lichenized symbiotic taiga association bryophytes foliose cyanobacteria ascomycetes lobaria tree genus russia microscopic trunk lichenology photosynthesis substratum ecosystem biome jagel