Determinants of bryophyte species composition and diversity on the Great Alvar of Öland, Sweden

Factors driving the species richness and distribution of bryophytes are poorly studied and not well understood, particularly in grasslands. We analysed the occurrence of bryophyte species and variation in species richness across 674 plots (0.5 m x 0.5 m) in alvar vegetation (grassland on limestone pavement with thin or no soil) on oland (Sweden) in relation to substrate characteristics and chemistry, inundation frequency, grazing pressure and geographical variables. We found 148 taxa, including 11 nationally red-listed ones. Species richness per plot was significantly associated with substrate type, positively associated with pH and grazing intensity, but negatively associated with soil depth. However, richness of species typical of, or restricted to, alvar habitats responded differently to richness of species more common in other habitats. Typical alvar species were favoured by high pH, shallow soil and low phosphate availability, while generalists preferred relatively low pH, higher phosphate availability and organic or mull soil types. Distance from the alvar margin had only weak effects. Concerning the effects on individual species and community composition, inundation frequency and pH were found to have the largest effects, although other factors (substrate type, soil depth, bare soil, bare stone, phosphate availability and grazing pressure) were more important for some individual species, stressing the importance of microsite variability and variability in management for regional species richness. From a conservation perspective, it is concluded that grazing is generally positive whilst factors increasing phosphate availability may disadvantage the typical alvar species, and proximity to the alvar margin is not a major problem.