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Heteronomous hyperparasitoids, sex ratios and adaptations: a reply

Donaldson & Walter (199la, b) described a series of observations and experiments designed to investigate the reproductive strategy of the he teronomous hyperparasitoid Coccophugus atratus (Hymenoptera: Aphelinidae). Heteronomous hyperparasitoids (also called autoparasitoids or adelphoparasitoids) are unusual in that male and female offspring develop obligatorily in different types of host. Donaldson & Walter found that sex ratio was strongly influenced by the proportion of hosts suitable for male and female offspring and strongly deviated from equality. They concluded that ‘interpretations based on [sex ratios] being adaptive seem inadequate’. We (Godfray & Hunter, 1992; see also Godfray & Waage, 1990) criticized the notion that a sex ratio of equality should always be expected to evolve in heteronomous hyperparasitoids. Specifically, we argued that observed sex ratios should be influenced by the factor limiting reproductive output. In circumstances where wasps search for both host types in the same environment, we considered the extreme cases of pure egg and pure time limitation. In the former case Fisher’s sex ratio theory applies and a 5050 sex ratio is predicted. However, in the second case, there is no longer a trade-off between the production of sons and daughters and the sex ratio will exactly reflect the relative proportions of hosts suitable for the two sexes. In the preceding short communication, Walter & Donaldson (1994) reply to our criticism. They accept our argument that Fisher’s prediction of a 50:50 sex ratio does not apply to host-limited heteronomous hyperparasitoids but (i) question whether our ideas apply to C.atrutu.s. (ii)