1 Zonation is often seen in environments with a strong physico-chemical gradient, such as salt marshes. It has been hypothesized that plant species are limited in their distribution by abiotic factors towards the more extreme end of the gradient, and by competition towards the more favourable end. Invasion of the native clonal grass genus Elymus in many Wadden Sea marshes may be due to increasing atmospheric nitrogen input into a nitrogen-limited environment. However, at Thmlauer Bay, Germany, Elymus athericus does not occur in lower salt marsh communities that are dominated by a dwarf shrub (Atriplex portulacoides). We therefore hypothesized that at this site the downslope (= more extreme) distributional boundary of E. athericus is a result of competition with A. portulacoides rather than of physiological limits.
2 A factorial experiment was set up to investigate the effects of removal of each competitor and fertilization. The reciprocal effects of the species on each other were measured in terms of vegetation cover and above-ground biomass. The impact of the tidal regime on plant zonation was investigated by calculating inundation frequencies at the boundary between the two plant populations from water level recordings.
3 Elymus athericus extended its distribution into the lower salt marsh when A. portulacoides was removed. The latter increased in cover but not in biomass after the removal of E. athericus. Neither species showed a response to nitrogen fertilization. The boundary between the two species in the control plots varied considerably in elevation and inundation frequency.
4 The lower distributional boundary of E. athericus can be interpreted as the result of competition with A. portulacoides. Improvement of nitrogen availability in concentrations of the same order of magnitude as annual atmospheric input had no detectable effect on plant zonation and growth. Elevation and inundation frequency were not strictly correlated with plant zonation.