Sex ratios and multifaceted parental investment

Although theories of parental investment and sex ratio generally assume that a single resource limits reproduction, many organisms invest two or more qualitatively different types of resources in the production of offspring. We examine the consequences of multifaceted parental investment for offspring provisioning and sex allocation, building our argument around a study of the nest-building Hymenoptera (wasps, bees, and ants). We review empirical studies that demonstrate that lifetime reproductive success may be constrained not only by resources used to provision offspring but also by the supply of mature oocytes or, in some cases, by the availability of space within nest sites or the time required to defend nests. Under multifaceted parental investment, the factor limiting parental fitness determines the currency of the optimization problem; parents are predicted to adjust reproductive behavior to maximize fitness returns per unit of the limiting resource. We develop simple models that predict that a greater availability of resources used for provisions will lead to an increase in the amount provisioned per offspring and an increase in the numerical or biomass proportion of females produced. These predictions explain widely observed patterns of variation in offspring provisioning and sex allocation in the nest-building Hymenoptera.