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1 The relationships between soil development, vertical vegetation zonation, vegetation succession and herbivory by Brent geese, Branta bernicla, were studied in a coastal salt marsh. We were able to analyse up to 100 years of salt marsh development by comparing sites where vegetation succession had progressed for various periods of time. These data were related to a continuous daily record of high water levels measured since 1824. 2 Most elevational variation in edaphic conditions (and therefore vertical vegetation zonation) could be attributed to variation in height of the sandy subsoil, as rapid dune formation occurs on the beaches early in succession. In the intermediate part of this elevational gradient, the maximum annual increase of 1.2 mm of silt corresponded to an annual increase of 5.6 g N [m.sup.-2] in the topsoil (0-50 cm). The average sea level rise in this area over the last 170 years was 0.63 mm [year.sup.-1]. A sedimentation model suggests that this has had strong effects on sedimentation and the annual inundation frequency in the mid-part of the elevational gradient, thus affecting vegetation zonation on the salt marsh. For the major part of the investigated transects, sea level rise has probably speeded up succession due to an increased rate of sedimentation. 3 The occurrence and dominance of all plant species were recorded in 3927 plots, and and for the 11 most common species response surfaces were calculated for their dependence on elevation and transect age. Most plant species were clearly separated along these axes. Most halophytic species, which were preferred by the geese, occurred early in succession and low on the gradient, where we observed the highest densities of Brent geese grazing. Forage quality of Festuca rubra increased towards the lower salt marsh. Other preferred forage species (Puccinellia maritima and Plantago maritima) were gradually displaced during succession by the tall grass Elymus athericus, especially in the mid- and upper salt marsh. Few geese grazed in areas where Elymus was dominant. 4 Herbivores first increased in numbers but then declined along a gradient of primary productivity. We propose that declining forage quality (due to changing vegetation composition during succession) is a better explanation for this pattern than the classic explanation of predator control of herbivores at high levels of primary productivity. This quality threshold hypothesis, as an alternative explanation of the exploitation ecosystem hypothesis, is expected to hold especially where smaller (quality-sensitive) herbivores such as geese are present. 5 Grazing by cattle in a 200-year-old part of the salt marsh led to the disappearance of Elymus athericus, to establishment of early successional halophytes and a return of Brent geese. Grazing by a larger herbivore therefore facilitated conditions for smaller herbivores by preventing the dominance of plant species that were good light competitors, and thus improved forage quality. Populations of these small herbivores could then become regulated by predators, although nine was present at our site. Keywords: facilitation, herbivory, nitrogen accumulation, salt marsh, sea level rise, succession

1 We analysed the germination of 91 herbaceous species in response to a temperature gradient, and to alternating temperatures and stratification (chilling). A principal-components analysis revealed that the species were distributed along two statistically independent axes, the first of which primary represented the optimal temperature for germination and the second the rate of germination. 2 These results were further related to data on species replacement during 25 years of succession in grassland after the cessation of fertilizer application but with continued hay making. Separate successional sequences were distinguished for the dry parts and the wet parts of the fields. Changes in canopy structure were studied by counting gaps and mapping light at the soil surface. 3 Changes in species composition of species present at the different stages were related to the changes in the germination attributes during the succession towards nutrient-poor grassland. The productive grassland was characterized by rapidly germinating species and in the wetter parts, by those germinating at low temperatures. This might enable these species to escape from light competition by germinating in the autumn or winter, i.e. soon after seed set. The species from the less productive, more open stages germinated more slowly, and responded more clearly to stratification and alternating temperatures. It is therefore likely that germination of most seeds of these species is delayed until the following summer. 4 The importance of the observed interspecific differences in germination characteristics in relation to the observed changes in vegetation structure as an explanation for species replacement during succession is discussed.

1 The present study reports on a primary succession series which started on bare soil on the Dutch island of Schiermonnikoog after the building of a sand dike. Vegetational changes were studied for 18 years by means of permanent transects along a topographic gradient from a moist plain to dry dunes. Soil development and vegetation structure were reconstructed using a chronosequence. A fertilizer experiment was set up in an intermediate successional stage in the plain and on the dune, in order to determine which soil resources limited productivity. 2 Differences in salinity, flooding and moisture content were important determinants of the differences in species composition along the topographic gradient. In addition, year-to-year fluctuations of these factors seem to be responsible for the year-to-year fluctuations in frequency of occurrence of many short-lived species. These factors did not, however, show a consistent long-term trend over time. 3 From soil analyses and the nutrient addition experiment, it is concluded that nitrogen limited above-ground biomass production. Over a period of about 16 years the total amount of nitrogen in the organic layer of the soil increased from 7 to 50 g N m to the -2 in the plains and from 1 to 15 g N m to the -2 on the dunes. 4 The accumulation of nitrogen during the successional series is accompanied by an increased biomass, a decreased light penetration to the soil surface, a decreased root/shoot ratio, increasing dominance of tall species, and a decreasing abundance of small, short-lived species. These data suggest that the importance of light competition is increasing during succession. 5 The importance of plant height versus light reduction at the soil surface in determining the outcome of light competition is discussed.