Evolution of size-dependent flowering in a variable environment: partitioning the effects of fluctuating selection

In a stochastic environment, two distinct processes, namely nonlinear averaging and non-equilibrium dynamics, influence fitness. We develop methods for decomposing the effects of temporal variation in demography into contributions from nonlinear averaging and non-equilibrium dynamics. We illustrate the approach using Carlina vulgaris, a monocarpic species in which recruitment, growth and survival all vary from year to year. In Carlina the absolute effect of temporal variation on the evolutionarily stable flowering strategy is substantial (ca. 50% of the evolutionarily stable flowering size) but the net effect is much smaller (ca. 10%) because the effects of temporal variation do not influence the evolutionarily stable strategy in the same direction.