Experimental separation of resource quantity from temporal variability: seedling responses to water pulses.

We tested the hypothesis that higher temporal variability in water supply will promote higher species richness of germinating and surviving seedlings using assemblages of 70 species of herbaceous plants from limestone pavement habitats. In a two-factor greenhouse experiment, doubling the total volume of water added led to greater germination (measured as number of germinated seeds and species) and establishment (survival and biomass) but the effects of temporal variability depended on the response variable considered. Low pulse frequencies of water addition with total volume added held constant resulted in greater temporal variability in soil moisture concentration that in turn promoted higher density and richness of germinated seedlings. Low pulse frequencies caused an eight-fold greater mortality in the low total volume treatment and biomass production to decline by one-third in the high total volume treatment. The effects of increasing temporal variability in water supply during recruitment stages can thus be opposite on different components of plant fitness and may also depend on total resource quantity. While greater species richness in more temporally variable soil moisture conditions was attributable to sampling effects rather than species-specific responses to the water treatments, species relative abundances did vary significantly with temporal variability. Changes in the amplitude or frequency of resource fluctuations may alter recruitment patterns, and could have severe and relatively rapid effects on community structure in unproductive ecosystems.