Persistent sex-by-environment effects on offspring fitness and sex-ratio adjustment in a wild bird population.

A major component of sex-allocation theory, the Trivers-Willard model (TWM), posits that sons and daughters are differentially affected by variation in the rearing environment. In many species, the amount of parental care received is expected to have differing effects on the fitness of males and females. When this occurs, the TWM predicts that selection should favour adjustment of the offspring sex ratio in relation to the expected fitness return from offspring. However, evidence for sex-by-environment effects is mixed, and little is known about the adaptive significance of producing either sex. Here, we test whether offspring sex ratios vary according to predictions of the TWM in the house wren (Troglodytes aedon, Vieillot). We also test the assumption of a sex-by-environment effect on offspring using two experiments, one in which we manipulated age differences among nestlings within broods, and another in which we held nestling age constant but manipulated brood size. As predicted, females with high investment ability overproduced sons relative to those with lower ability. Males were also overproduced early within breeding seasons. In our experiments, the body mass of sons was more strongly affected by the sibling-competitive environment and resource availability than that of daughters: males grew heavier than females when reared in good conditions but were lighter than females when in poor conditions. Parents rearing broods with 1:1 sex ratios were more productive than parents rearing broods biased more strongly towards sons or daughters, suggesting that selection favours the production of mixed-sex broods. However, differences in the condition of offspring as neonates persisted to adulthood, and their reproductive success as adults varied with the body mass of sons, but not daughters, prior to independence from parental care. Thus, selection should favour slight but predictable variations in the sex ratio in relation to the quality of offspring that parents are able to produce. Offspring sex interacts with the neonatal environment to influence offspring fitness, thus favouring sex-ratio adjustment by parents. However, increased sensitivity of males to environmental conditions, such as sibling rivalry and resource availability, reduces the fitness returns from highly male-biased broods.