Bryophyte species richness on boulders: relationship to area, habitat diversity and canopy tree species

Bryophytes growing on siliceous boulders (diameter 50-200 cm) were studied in two forests in east-central Sweden to investigate the influence of different canopy tree species on the bryophyte species richness. Granite boulders lying below crowns of Ulmus glabra (elm), Fraxinus excelsior (ash), Acer platanoides (maple), Quercus robur (oak), Betula pendula (birch), and Picea abies (Norway spruce) were studied in two mixed stands. Both boulder area and within-boulder habitat diversity were positively related to species richness. Boulders below the base-rich deciduous trees F. excelsior, U. glabra and A. platanoides held roughly twice as many species as those below P. abies, with Q. robur and B. pendula as intermediates. The rank order among trees was as expected from bark pH and litter decomposability in the literature. We also used sample plots to investigate species richness at a smaller scale. Within plots, intermediate levels of litter, inclination and exposed rock promoted species richness, and also the covering tree species had an effect. Individual plots were on average not more species-rich on large boulders than on small ones. This leads us to conclude that population extinctions on the plot-level, and re-colonisations from other parts of the boulder, are less important for species richness than within-boulder habitat diversity. Since species richness was lower under P. abies it is important to favour a diversity of tree species and to include base-rich deciduous trees in otherwise homogeneous Picea forests.