Competition among the pioneers in a seasonal soft-bottom benthic succession: field experiments and analysis of the Gilpin-Ayala competition model.

Controlled experiments, designed to assess the effects of pioneers on succession on an intertidal sandflat, provided evidence for interspecific competition between juvenile Hobsonia florida (Polychaeta, Ampharetidae) and oligochaetes. The field data were fitted to both the linear Volterra and non-linear Gilpin-Ayala competition equations. With its greater number of parameters, the Gilpin-Ayala model must provide a better fit to observed population abundances. The Gilpin-Ayala model is flawed as an explanation of the population trajectories of the H. florida and oligochaetes, because its non-linearity parameter affects only intraspecific competion. With either model our field data demonstrate a solution to Hutchinson's paradox. With competition coefficients near unity and similar carrying capacities, the predicted population trajectories are heavily dependent on initial conditions. The predicted times to competitive exclusion are long and can easily exceed the typical period of environmental constancy. Our study offers evidence for Neill's competitive bottleneck: competition acts primarily on the developmental stages of one of a pair of competing species. The permanent meiofauna may act as a competitive bottleneck for the population growth of benthic macrofauna. The mechanism of this competitive interaction probably involves exploitative interspecific competition for benthic diatoms.