Insights from complete-incomplete brood sex-ratio disparity

An assertion deeply rooted in the ornithological literature holds that sex-specific mortality causes a sex ratio disparity (SRD) between complete and incomplete broods. Complete broods are thought to reflect the primary sex ratio before any bias introduced by developmental mortality. Contrary to this view, however, complete and incomplete broods should exhibit identical sex ratio distributions even when the sexes experience differential mortality, as shown in the classic paper of Fiala (Am Nat 115: 442–444, 1980). Therefore, in partially unsexed samples, primary sex ratio biases cannot be distinguished from biases caused by differential mortality. In addition, complete broods do not represent primary sex ratio more accurately than incomplete ones and might even be misleading. Despite Fiala’s prediction, SRD does occur in some empirical studies. We show that this pattern could arise if (1) primary sex ratio affects chick mortality rates independently of sex (direct effect), (2) primary sex ratio covaries with a variable that also affects mortality rate, or (3) sex differential mortality covaries with overall mortality rate (indirect effects). Direct effects may cause stronger SRD than indirect ones with a smaller and opposite bias in the overall sex ratio and could also lead to highly inconsistent covariate effects on brood sex ratios. These features may help differentiate direct from indirect effects. Most interestingly, differences in covariate effects between complete and incomplete broods imply that influential variables are missing from the analysis.