The vegetation of coastal sand dunes is characterized by high species diversity and comprises some of the rarest vegetation types in North-Western Europe. Among them are dune grassland communities whose species richness relies on grazing. Those communities are assessed as a priority habitat type under the Natura 2000 legislation. In autumn 1990, Galloway cows and Nordic Fjord horses were introduced in the coastal dunes of Meijendel near The Hague (52°7'N, 4°20'E), The Netherlands, to reduce encroachment of tall grasses and shrubs, to develop bare sand patches, and as such facilitating diverse vegetation structures in the dune grasslands. In the 1950s, decades before the introduction of livestock, 41 permanent plots were installed. On average, they were examined every four years. Our study hypothesised that the livestock grazing in the set densities would halt progressive succession and facilitate regressive succession. Up to 1990, we observed an equilibrium between progressive and regressive succession. After 1990, however, our data showed a pronounced progressive succession contradicting the hypothesized effect of the livestock grazing. We relate the main observed patterns with two factors linked to rabbit populations: (i) the myxomatosis outbreak in 1954 and (ii) the rabbit Viral Haemorrhagic Disease (rVHD-1) outbreak in 1989. In addition to livestock grazing, rabbits block progressive succession by feeding on seedlings of shrub and tree species and digging burrows, creating small-scale mosaics of bare sand and initiate blowout development when collapsing. We state that the substantial decrease in rabbit numbers due to the viral diseases likely caused the observed increase of shrubs and trees in the study area's permanent plots. Climate change might have contributed to the observed increase in autonomous blowout development since 2001, as well as a decrease in atmospheric nitrogen deposition since 1990, after a strong increase the decades before.
