Born without a Silver Spoon: A Review of the Causes and Consequences of Adversity during Early Life

A huge amount of research attention has focused on the evolution of life histories, but most research focuses on dominant individuals that acquire a disproportionate level of reproductive success, while the life histories and reproductive tactics of subordinate individuals have received less attention. Here, we review the links between early life adversity and performance during adulthood in birds, and highlight instances in which subordinate individuals outperform dominant conspecifics. Subordinate individuals are those from broods raised under high risk of predation, with low availability of food, and/or with many parasites. Meanwhile, the broods of many species hatch or are born asynchronously and mitigation of the asynchrony is generally lacking from variation in maternal effects such as egg size and hormone deposition or genetic effects such as offspring sex or parentage. Subordinate individuals employ patterns of differential growth to attempt to mitigate the adversity they experience during early life, yet they overwhelmingly fail to overcome their initial handicap. In terms of surviving through to adulthood, subordinate individuals employ other "suboptimal" tactics, such as adaptively timing foraging behaviors to avoid dominant individuals. During adulthood, meanwhile, subordinate individuals rely on "suboptimal" tactics, such as adaptive dispersal behaviors and competing for partners at optimal times, because they represent the best options available to them to acquire copulations whenever possible. We conclude that there is a gap in knowledge for direct links between early life adversity and subordination during adulthood, meaning that further research should test for links. There are instances, however, where subordinate individuals employ "suboptimal" tactics that allow them to outperform dominant conspecifics during adulthood.