Sex allocation theory has provided a remarkably successful framework for predicting investment in offspring sex or in egg versus sperm for invertebrates and plants (Charnov 1982;Wrensch and Ebbert 1993;Hardy 2002), yet sex ratio biases in amniote vertebrates seem unsystematic and often are explained a posteriori. The frequent mismatch between theory and empirical data in amniotes raises doubts as to whether adaptive sex allocation occurs in these taxa and highlights the need for expanded study focusing on long-lived organisms.Many hypotheses of sex allocation for amniote vertebrates are based on condition dependence of sex allocation (Charnov 1982; Cockburn et al. 2002;West 2009). An individual invests in sons versus daughters or eggs versus sperm depending on individual condition, such as resource availability, body size, or incubation temperature. That a sex-specific fitness advantage of body size might select for size-dependent sex allocation was first proposed for sex-changing invertebrates (Ghiselin 1969). Subsequently, the Trivers-Willard hypothesis (TW; Trivers and Willard 1973) proposed that offspring sex in mammals may be linked to maternal condition if condition influences the lifetime fitness of one offspring sex more than the other. Charnov and Bull (1977) (see also Bull 1981) expanded condition-dependent sex allocation to environmental sex determination, providing an adaptive hypothesis for the occurrence of temperature-dependent sex determination (TSD) that proposes developmental temperature is the fitness-altering condition that determines investment in ovaries or testes (Gutzke and Crews 1988; Janzen 1995;Shine 1999;Warner and Shine 2008). Condition dependence in sex allocation has been expanded to sex biases based on individual size, paternal quality, parental rank, hatching order, and season (Werren and Charnov 1978;Charnov 1979a;Burley 1981;Silk 1983;Bednarz and Hayden 1991;Daan et al. 1996).According to sex allocation theory, sex allocation should be based on individual condition relative to the condition of competitors, not on absolute condition (Charnov et al. , 1981 Charnov 1982;West 2009). The TW hypothesis predicts that mothers of relatively good condition should invest more in sons than mothers of relatively poor condition (Trivers and Willard 1973). This is because each sex competes for lifetime reproductive success only with members of the same sex, so that the quality of a son or daughter is defined only with respect to their sexual competitors (other sons and daughters in the population). If sex allocation were based on absolute condition, a year with extreme condition values would vastly overproduce one sex, and frequency-dependent selection on sex would favor a mutant that produced the rare sex. The resulting prediction is that the optimal sex allocation versus condition reaction norm should be relative, shifting in response to changes in the population distribution of condition, and that an equilibrium cohort sex ratio would be maintained. When individuals are not allowed to def...