Juniper Shade Enables Terricolous Lichens and Mosses to Maintain High Photochemical Efficiency in a Semiarid Temperate Sand Grassland

On a semiarid sand grassland (Festucetum vaginatae) colonised by juniper (Juniperus communis L.) shrubs terricolous lichens and mosses segregate strongly between microhabitats: certain species grow in the open grassland, others almost exclusively in the shade of junipers. The contrasting irradiances of these microhabitats influence much the metabolism of these organisms, and thus affect their small-scale distribution. This was confirmed by determining the efficiency of photochemical energy conversion by measuring chlorophyll a fluorescence parameters. In the open grassland maximum photochemical efficiency of photosystem 2 (PS2, F_v/F_m) declined from the humid spring to the hot and dry summer in all species, and this was caused by an increase in base fluorescence (F₀), but not by the decrease in fluorescence maximum (F_m). In summer, mosses and lichens growing in the open grassland generally possessed lower F_v/F_m than cryptogams growing in the shade cast by juniper shrubs. Thus mosses and lichens in the open grassland suffer lasting reduction in photochemical efficiency in summer, which is avoided in the shade of junipers. Juniper shrubs indeed influence the composition and small-scale spatial pattern of sympatric terricolous lichen and moss communities by—among others—providing a shelter against high light in summer.