Sex expression and genotypic sex ratio vary with region and environment in the wetland moss Drepanocladus lycopodioides

Abstract

Sex ratio variation is common among organisms with separate sexes. In bryophytes, sex chromosome segregation at meiosis suggests a balanced progeny sex ratio. However, most bryophyte populations exhibit female-biased phenotypic sex ratios based on the presence of reproductive structures on gametophytes. Many bryophyte populations do not form sexual organs, and genotypic sex ratio variation in such populations is mostly unknown. We tested sex expression, and phenotypic and genotypic sex ratios against environmental parameters in natural populations of the unisexual wetland moss Drepanocladus lycopodiodes at 11 sites in each of three regions in southern Sweden. We identified sex in 660 individual ramets, based on sexual structures, when present, or with a specifically designed molecular marker, when absent. All regions exhibited a female bias in phenotypic and genotypic sex ratios. Sex ratio biases and sex expression differed between regions. Sex ratios were less female-biased in larger patches. Wetter patches exhibited a stronger female bias in genotypic sex ratio and lower sex expression. This is the first evidence of environmental effects on genotypic sex ratio in mosses. A higher frequency of females in wet patches could be due to higher female resource demands for sporophyte production or higher male sensitivity to wetness. A higher incidence of females than males in moister sites aligns with female flowering plants, but differs from reproductive bryophytes in drier environments. Taken together with previous results, our data indicate that sex ratio variation and its drivers differ among species, their life histories and environments.