INTRAPOPULATION DIFFERENTIATION IN ANNUAL PLANTS. III. THE CONTRASTING EFFECTS OF INTRA- AND INTERSPECIFIC COMPETITION.

The roles of intraspecific and interspecific competition in producing differentiation within populations of Veronica peregrina were studied in two populations under controlled, greenhouse conditions. In nature, each population spans an environmental gradient across the center and sides of a temporary, vernal pool in California. Individuals at the center are subjected to intense intraspecific competition produced by high densities (to 30 seedlings/cm ) generated by quasi-simultaneous germination (90% of seeds germinate in one week). Individuals at the periphery are subjected to interspecific competition with grasses, which shade out the Veronica 4-6 weeks after the onset of winter growth. I predicted that 1) when grown under immediate intraspecific competition in the greenhouse, offspring of plants from the central subpopulation (C) would perform better (i.e., grow larger and produce more seeds) than those from the periphery (P) and that 2) when grown under delayed interspecific competition provided by Agrostis tenuis and Lollium multiflorum, offspring of plants from the periphery would perform better than those from the center. Both predictions were confirmed. The center-periphery differences were pronounced and statistically significant in an undisturbed population (V-2), while in a population disturbed by yearly plowing (V-3), the differences tended to be consistent with those in V-2 but seldom significant. Distribution of variability tended to be positively skewed and/or leptokurtic in subpopulations grown under "foreign" competition (i.e., intraspecific for P plants and interspecific for C plants) but was normally distributed following exposure to "familiar" competition. Timing of competition affected many results. There were four additional significant differences between the central and peripheral subpopulations. 1) Germination rate: the faster rate in central plants can be advantageous under immediate intraspecific competition. The slower rate in peripheral plants can be advantageous under conditions of erratic and unpredictable soil moisture. 2) Response to nutrient competition: central plants were more sensitive to N-deficiency and peripheral plants were more sensitive to P-deficiency. 3) Allocation of biomass: central plants allocated a greater proportion of biomass to seeds, while peripheral plants allocated a greater proportion of biomass to leaves under all growing conditions. 4) Root elongation: at the seedling stage, central plants have longer roots, while at the adult stage, peripheral plants have longer roots (but not more root mass). Most components of this complex pattern of differentiation are interpretable in an adaptive context. Other results defy simple explanations and underline the importance of phenotypic plasticity, which was pronounced in the competition experiments.