Effects of Long-Term Trampling on the Above-Ground Forest Vegetation and Soil Seed Bank at the Base of Limestone Cliffs

Exposed limestone cliffs in central Europe harbor a highly divers flora with many rare and endangered species. During the past few decades, there has been increasing recreational use of these cliffs, which has caused local environmental disturbances. Successful restoration strategies hinge on identifying critical limitations. We examined the composition of aboveground forest vegetation and density and species composition of seeds in the soil seed bank at the base of four limestone cliffs in mixed deciduous forests that are intensively disturbed by human trampling and at four undisturbed cliffs in the Jura Mountains in northwestern Switzerland. We found that long-term human trampling reduced total aboveground vegetation cover at the base of cliffs and caused a significant shift in the plant-species composition. Compared with undisturbed cliffs, total seed density was lower in disturbed cliffs. Human trampling also altered the species composition of seeds in the soil seed bank. Seeds of unintentionally introduced, stress-tolerant, and ruderal species dominated the soil seed bank at the base of disturbed cliffs. Our findings indicate that a restoration of degraded cliff bases from the existing soil seed bank would result in a substantial change of the original unique plant composition. Active seed transfer, or seed flux from adjacent undisturbed forest areas, is essential for restoration success.