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400 algae ascomata atmospheric colonizer colonizers colorful common composite conditions crustose cyanobacteria dehydration diaspores digestion early ecosystem encrusting evolution external extreme filaments forming fungal fungi fungus growing growth habitat harsh lichen living longlived maritime medicinal million moss mutualistic natural old oldest orange organism organisms park patch photosynthesis pigments plant pollution rate relationship reproduction sensitive shore slow sunburst symbiosis terrestrial texture thallus tree vegetative xanthoria years yellow