Abstract
Contrary to our expectations, soil salinity and moisture explained little of the spatial variation in plant establishment in the upper intertidal marsh of three southern California wetlands, but did explain the timing of germination. Seedlings of 27 species were identified in 1996 and 1997. The seedlings were abundant (maximum densities of 2143/m2 in 1996 and 1819/m2 in 1997) and predominantly annual species. CCAs quantified the spatial variation in seedling density that could be explained by three groups of predictor variables: (1) perennial plant cover, elevation and soil texture (16% of variation), (2) wetland identity (14% of variation) and (3) surface soil salinity and moisture (2% of variation). Increasing the spatial scale of analysis changed the variables that best predicted patterns of species densities. Timing of germination depended on surface soil salinity and, to a lesser extent, soil moisture. Germination occurred after salinity had dropped below a threshold or, in some cases, after moisture had increased above a critical level. Between 32% and 92% of the seedlings were exotic and most of these occurred at lower soil salinity than native species. However, Parapholis incurva and Mesembryanthemum nodiflorum were found in the same environments as the native species. In 1997, the year of a strong El Nino/Southem Oscillation event with high rainfall and sea levels, the elevation distribution of species narrowed and densities of P. incurva and other exotic species decreased but densities of native and rare species did not change. The 'regeneration niche' of wetland plant communities includes the effects of multiple abiotic and biotic factors on both the spatial and temporal variations in plant establishment.
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