Seed origin determines the range expansion of the clonal grass Elymus athericus

The recent invasion of clonal grasses to novel habitats poses a threat to biodiversity in various habitats. Elymus athericus, a clonal grass of north-western European salt marshes, is currently increasing in abundance and invading new habitats. In this study, we analyzed controlling factors for seedling establishment of E. athericus in frequently flooded low marsh habitats. Here, biotic and abiotic conditions are very different from the conditions of the parental sites with established populations higher up in the marsh. Hence, we hypothesized that seedling establishment at the expanding low marsh edge would depend on the parental origin (either through maternal effects or heritable local adaptation). We further hypothesized that seedling origin interacts with biotic factors such as herbivory and competition as well as with abiotic factors like inundation frequency. We tested the dependence of seedling survival, growth and vegetative reproduction on these factors in a factorial transplant experiment on Schiermonnikoog. Survival was high, with 77% of the planted seedling surviving until the end of the experiment. Biotic factors had a much stronger effect on seedling growth and mortality than parental origin and were independent of inundation. However, parental origin strongly interacted with herbivory and competition, with seedlings performing better under the conditions that resembled their parental site.We conclude that seedlings of E. athericus, a species that was previously thought to occur only in mid- to high marsh elevation, can establish at a frequently inundated low-marsh sites. Long term survival and further invasion will primarily depend on biotic factors in interaction with seed origin. Our results suggest that next to herbivory, limitation of seeds adapted to colonizing conditions is likely to slow down range expansion.