Co-evolution of seed size and seed predation

Using the evolutionarily stable strategy (ESS) approach in a model for the co-evolution of seed size and seed predation, I show that seed size variation within individual plants is favoured if there is a trade-off in the predator's attack rate for different seed sizes. A single seed size is not evolutionarily stable because a predator that is optimally adapted to one particular seed size cannot prevent invasion by plants with a different seed size. The model generates the following predictions. The ESS consists of a continuous range of seed sizes. Small seeds tend to be attacked more frequently than big seeds. Plants with many resources and plants with low (frequency-independent) juvenile mortality have more variable seeds than plants with few resources and a high juvenile mortality. Seed size variation is higher in fluctuating populations regulated by seed predation alone than in stable populations (partially) regulated by seedling competition. Predator searching behaviour does not directly affect the ESS seed size range, but may have an indirect effect by affecting population stability or the significance of seedling competition as a population regulating mechanism. Moreover, seed size distributions are found to be more skewed in favour of small seeds if predation is spatially non-uniform than if predation is more even. Application of the model to systems of several co-evolving plant and predator species is discussed.