Maternal effects and range expansion: a key factor in a dynamic process?

Species that depend on ephemeral habitat often evolve distinct dispersal strategies in which the propensity to disperse is closely integrated with a suite of morphological, behavioural and physiological traits that influence colonizing ability. These strategies are maintained by natural selection resulting from spatial and temporal variation in resource abundance and are particularly evident during range expansion. Yet the mechanisms that maintain close alignment of such strategies with resource availability, integrate suites of dispersal traits and generate variability in dispersal propensity are rarely known. Breeding females can influence offspring phenotype in response to changes in current environmental conditions, making maternal effects uniquely suited to bridge fluctuations in resource abundance in the maternal generation and variation in offspring dispersal ability. Western bluebirds' (Sialia mexicana) dependence on nest cavities--an ephemeral resource--has led to the evolution of two distinct dispersal phenotypes: aggressive males that disperse and non-aggressive males that remain philopatric and cooperate with their relatives. Over the last 40 years, western bluebirds rapidly expanded their geographical range, providing us with an opportunity to test, in newly established populations, the importance of maternal effects for generating variability in dispersal propensity. Here, I show that, under variable resource conditions, breeding females group offspring of different competitive ability in different positions in the egg-laying order and, consequently, produce aggressive males that are more likely to disperse when resources are low and non-aggressive philopatric males when resources are abundant. I then show experimentally that the association between resource availability and sex-specific birth order is robust across populations. Thus, this maternal effect enables close tracking of resource availability and may explain how variation in dispersal is generated in newly colonized populations. More generally, these results suggest that, as a key source of variation in colonizing phenotypes, maternal effects are of crucial importance for understanding the dynamics of range expansion.