Coping with environmental uncertainty: dynamic bet hedging as a maternal effect.

Mothers in a range of taxa manipulate the phenotype of their offspring in response to environmental change in order to maximize their own fitness. Most studies have focused on changes in the mean phenotype of offspring. Focusing on mean offspring phenotypes is appropriate for species in which mothers are likely to successfully predict the environment their offspring will experience, but what happens when the offspring's environment is unpredictable? Theory suggests that when mothers face uncertainty regarding their offspring's environment, they should increase within-clutch variation in the offspring phenotype (i.e. they should bet hedge). While comparative analyses support the idea that mothers do bet hedge in response to environmental unpredictability, empirical tests are very rare and it remains unclear whether mothers adaptively adjust variance in offspring traits (a phenomenon we call dynamic bet hedging). As a first step towards examining dynamic bet hedging, we reanalysed data from five previously published studies. These studies were across a range of taxa, but all manipulated the maternal environment/phenotype and then examined changes in mean offspring size. We found some support for the theoretical predictions that mothers should increase within-clutch offspring size variation when faced with unpredictable environments. We predict that dynamic bet hedging is more common than previously anticipated and suggest that it has some interesting implications for the studies that focus on shifts in mean offspring traits alone. Hence, future studies should examine maternal effects on both the mean and the variance of offspring traits.