Spiteful soldiers and sex ratio conflict in polyembryonic parasitoid wasps

The existence of spiteful behaviors remains controversial. Spiteful behaviors are those that are harmful to both the actor and the recipient, and they represent one of the four fundamental types of social behavior (alongside selfishness, altruism, and mutual benefit). It has generally been assumed that the conditions required for spite to evolve are too restrictive, and so spite is unlikely to be important. This idea has been challenged in recent years, with the realization that localized competition can relax the conditions required for spite to evolve. Here we develop a theoretical model for a prime candidate for a spiteful behavior, the production of the sterile soldier caste in polyembryonic wasps. Our results show that (a) the biology of these soldiers is consistent with their main role being to mediate conflict over the sex ratio and not to defend against competitors and (b) greater conflict will occur in more outbred populations. We also show that the production of the sterile soldier caste can be classed as a spiteful behavior but that, to an extent, this is merely a semantic choice, and other interpretations such as altruism or indirect altruism are valid. However, the spite interpretation is useful in that it can lead to a more natural interpretation of relatedness and facilitate the classification of behaviors in a way that emphasizes biologically interesting differences that can be empirically tested.