Long-Term Effects of Sheep Grazing in Various Densities on Marsh Properties and Vegetation Dynamics in Two Different Salt-Marsh Zones

For conservation management of grassland ecosystems, an important question is under which conditions large grazers induce compositional and structural variation in plant communities, which is a prerequisite for high biodiversity. Here we used two long-term projects on the mainland salt marshes of the Wadden Sea to test the hypothesis that long-term grazing management with different stocking densities results in plant communities with distinctively different plant species composition and vegetation structure. The two projects took place on a low clayey and a high sandy salt marsh with different stocking densities of sheep: 0, 1.5, 3.5, 4.5 and the initially 10 sheep ha⁻¹, where measurements were collected 11, 15, 19 and 23 years after the start of the project. Moreover, grazers affect abiotic conditions by reducing soil-redox potential and surface elevation, thereby driving composition and structure of salt-marsh vegetation. On the low salt marsh, a continued high stocking density (10 sheep ha⁻¹) resulted in succession from the early-successional Puccinellia maritima community to the late-successional Atriplex portulacoides community. On the high salt marsh, the early-successional Festuca rubra community was maintained under all stocking densities. Cessation of grazing resulted in succession to the Elytrigia atherica community in both salt-marsh types. Intermediate stocking densities (1.5, 3 or 4.5 sheep ha⁻¹) resulted in a mosaic of tall vegetation and patches of lawn, i.e. short-grazed vegetation, where Puccinellia maritima lawn occurred interspersed with patches of the Festuca rubra and tall Elytrigia atherica communities in both salt-marsh types. Effects of grazers were influenced by the presence of watering points near the sea wall. To conclude, our results show how joint interactions between grazers and abiotic conditions drive vegetation diversity and heterogeneity, with implications for ecosystem functions and services such as wildlife biodiversity and coastal protection.