Size and the Not-So-Single Sex: Disentangling the Effects of Size and Budget on Sex Allocation in Hermaphrodites.

Sex allocation theory explains how size-related variations in male and female fitness may favor the evolution of size-dependent sex allocation in hermaphrodites. Although empirical studies show that sex allocation changes gradually with size in many species, theoretical studies tend to predict an abrupt sex reversal from one sex to the other, that is, single-sexed sequential hermaphrodites. We show that this discrepancy between data and theory collapses if one takes into account that size affects male and female fitness through distinct routes. Using the classification of budget (larger individuals spend a greater budget on reproduction) and direct (e.g., larger plants are taller and may disperse pollen more efficiently) effects of size suggested by Klinkhamer et al., we propose a simple general framework appropriately incorporating these two categories of size effects in male and female fitness expressions. Analytical and numerical results show that a gradual sex change is evolutionarily stable for a large set of parameter values. Sex reversal is selected only in the absence of budget effects of size. We provide further predictions on size-dependent sex allocation and assess the relative importance of budget and direct effect for creating different patterns.