Abstract
To understand colonization processes, it is critical to fully assess the role of dispersal in shaping biogeographical patterns at the gene, individual, population, and community levels. We test two alternative hypotheses (H I and H II) for the colonization of disturbed sites by clonal plants, by analyzing intraspecific genetic variation in one and reproductive traits in two typical fen mosses with separate sexes and intermittent spore dispersal, comparing disturbed, early-succession (limed) fens and late-successional rich fens. H I suggests initial colonization of disturbed sites by diverse genotypes of which fewer remain in late-successional fens and an initially balanced sex ratio that develops into a possibly skewed population sex ratio. H II suggests initial colonization by few genotypes and gradual accumulation of additional genotypes and an initially skewed sex ratio that alters into the species-specific sex ratio, during succession. Under both scenarios, we expect enhanced sexual reproduction in late-successional fens due to resource gains and decreased intermate distances when clones expand. We show that the intraspecific genetic diversity, assessed by two molecular markers, in Scorpidium cossonii was higher and the genetic variation among sites was smaller in disturbed than late-successional rich fens. Sex ratio was balanced in S. cossonii and Campylium stellatum in disturbed fens and skewed in C. stellatum in late-successional fens, thus supporting H I. In line with our prediction, sex expression incidence was higher in, and sporophytes were confined to, late-succession compared to disturbed rich fens. Late-successional S. cossonii sites had more within-site patches with two or more genotypes, and both species displayed higher sex expression levels in late-successional than in disturbed sites. We conclude that diverse genotypes and both sexes disperse efficiently to, and successfully colonize new sites, while patterns of genetic variation and sexual reproduction in late-successional rich fens are gradually shaped by local conditions and interactions over extended time periods.