Bryophytes in black alder swamps in south Sweden: habitat classification, environmental factors and life-strategies

A considerable part of black alder swamps in Europe are situated in south Sweden. These swamps are frequently rich in bryophytes, but the variation in species composition is often large between sites. In this study we focus on environmental variables that best predict biodiversity in terms of species number, cover and bryophyte composition. The bryophyte vegetation was analysed in 71 plots of size 10 × 10 m in 31 black alder swamps from south Sweden. A total number of 171 bryophyte species (including 52 liverwort species) were recorded. At plot level the species number ranged between 11 and 54 (0–19 for liverworts and 11–36 for mosses). The black alder swamps were classified into five groups, which turned out to be characterized by a combination of tree species composition and hydrological conditions. 22 environmental and geographic variables were recorded for each plot and used as predictor variables in a redundancy analysis (RDA). These variables explained about 50% of the distribution of bryophytes. The most influential variables along the first axis were slope (positive relationship), basal area of spruce (positive relationship) and basal area of black alder (negative relationship). Along the second axis slope, basal area of ash and water depth had the strongest impact. The variables were subdivided into four groups and analyzed by separate RDAs. The amount of dead wood was important for the species richness in black alder fens in combination with humidity. Epixylic bryophytes tended to occupy spruce dead wood as a primary substrate and alder dead wood as a secondary substrate if spruce is present. Black alder stools were important as substrate for various bryophytes during flooding periods, including Plagiothecium latebricola and some liverworts (Calypogeia spp. and Riccardia spp.). The life-strategy categories were represented by perennial stayers 11%, competitive perennials 28%, stress-tolerant perennials 9%, colonists sensu stricto 21%, pioneer colonists 13%, long-lived shuttles 15% and short-lived shuttles 3%. The composition of life-strategy types varied in the swamps. Stress-tolerant perennials were more common in flat swamps. Colonists were more frequent in swamps with dominance of spruce and in sloping swamps