THE GENETIC-BASIS OF PHOTOPERIODISM AND ITS EVOLUTIONARY DIVERGENCE AMONG POPULATIONS OF THE PITCHER-PLANT MOSQUITO, WYEOMYIA-SMITHII

We measured the additive genetic variance within populations and the composite additive, dominance, and epistatic effects contributing to differentiation of photoperiodic response between two southern (ancestral) and each of four progressively more northern (derived) populations of the pitcher-plant mosquito, Wyeomyia smithii. Critical photoperiod and its additive genetic variance but not its heritability increased with latitude. Directional selection on critical photoperiod during the northward divergence of W. smithii has therefore not eroded the additive genetic variance underlying this trait. Joint scaling tests of crosses between populations showed that epistatic effects, especially additive x additive and dominance x dominance interactions, overwhelm composite additive and dominance effects on critical photoperiod. The presence of substantial epistasis suggests that multiple founder events during the northward divergence of W. smithii may have been responsible for the release of progressively greater additive genetic variance in derived populations, despite directional and stabilizing selection to reduce it. If epistasis makes a similar contribution to the genetic differentiation of populations in other species, then current models of adaptive evolution that consider only additive genetic variation and covariation within populations may be of limited value in predicting how natural populations differentiate in life history.