The Trivers-Willard hypothesis (TWH) states that parents in good condition preferentially produce the sex with a higher variation in reproductive success, whereas parents in bad condition favour the opposite sex. Theorists distinguish two variants of the TWH: (a) a biased sex-ratio at birth and (b) biased parental investment after birth. It has been argued before that the conditions stated by Trivers and Willard (good condition is inherited and affects reproductive success more strongly for one of the sexes) are sufficient for the sex-ratio version but insufficient for the investment version of the TWH. However, it has not yet been investigated how these conditions affect parental investment in high and low quality parents, depending on the life-cycle of a species. The present study aims to fill this gap by introducing a multi-stage matrix population model with nonlinear mating to describe the effects of parental investment after birth on the reproductive values of male and female individuals. Using methods from adaptive dynamics and evolutionary invasion analysis, evolutionary trajectories and evolutionarily stable strategies are derived for different parameterizations of the model. Simulation results demonstrate that the conditions given by Trivers and Willard produce a general bias of parental investment towards the sex with higher variance in reproductive value. This bias is stronger for low-quality parents than for high-quality parents and matches the expected marginal offspring reproductive values for parental investment.
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