Some conceptual issues in the origin of eusociality

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Abstract
Certain issues arising in connection with the evolutionary origins of eusociality are discussed. Previous results about when natural selection favours helping behaviour are generlised, and the differing viewpoints of both parents and offspring are considered. Particular attention is given to the evolutionary implications of different patterns of overlapping generations observed in bivoltine insects. As argued by Seger (1983), these patterns imply different conditions under which a daughter is selected to help her mother rear additional siblings in haplodiploid populations. Other factors that can alter the selective advantages of helping behaviour under haplodiploidy are also discussed, including the possibility of sex ratio manipulation and the novel result that helping behaviour may be locally favoured in populations that are spatially patchy with respect to sex-specific fitness. A new hypothesis is also presented: The fact that sisters are selected to aid their mother to parasitise other sisters may have played an important role in the origins of eusociality. A given offspring benefits from having maternally parasitised siblings because such siblings rear additional siblings (to which the given offspring is more closely related) instead of nieces and nephews. Finally, the importance of haploidiploidy in the origins of eusociality is discounted; the virtually unique biology of aculeate Hymenoptera would seem to be of much greater importance.