Population biology of salt marsh and sand dune annuals

Annuals represent a significant component of the vegetation of coastal salt marshes and sand dunes. From many points of view, the two habitats might appear to have little in common. Yet both are characterized by episodes of low water potential, marked spatial and temporal heterogeneity and a zonation which, within certain limits, reflects successional change. There are also similarities of distribution. Annuals are dominant usually in the pioneer stages; the Salicornia-dominated low marsh areas are perhaps analogues with strandline ephemeral populations (e.g. Cakile maritima) on the fore-dunes. In mature stages, annuals are associated with small gaps in the matrix of perennials, at least some of these arising from drought or disturbance. Nevertheless populations can reach very high densities. The most striking contrast is phenological; only summer annuals are found on marshes, whereas winter annuals predominate on dunes (except for the strandline). Similarly there is a difference in species richness. Rather few species of annual are typical of marshes while a great many are found on dunes. Properties of the seed bank, survival, reproduction and population regulation are compared in marsh and dune annuals, with special reference to Cakile, Salicornia, Rhinanthus and Vulpia. Interpretations are suggested which take account of environmental predictability and heterogeneity. Finally, the general applicability of simple mathematical models of these populations in the different coastal habitats is considered.