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Living things are complex, but this complexity is of two broad types. Organisms show complexity of apparent purpose, with all of the parts acting for the whole. Groups, however, are usually dominated by the complexities of cross-purpose; the parts seem goal-directed, but the goals are not shared, and the result is often anything but elegant. The most spectacular exceptions, at the group level, are social insect societies, in which the individuals usually do seem to act toward a common goal.

Any scientific theory purporting to account for biological complexity ought to account for this special nature of social insects. Why do their colonies show a degree of apparent purpose lacking in other aggregations, herds, and flocks? The kin selection extension of natural selection theory does explain this; cooperation results from the opportunity to give sufficiently large benefits to kin.

More importantly, kin selection theory has successfully predicted new findings. Although social insect colonies have clock-like design in many respects, kin selection theory predicts who is throwing sand into the clockworks, as well as which gears might be slipped and which springs sprung. Many of the predicted findings, such as sex ratio conflict and policing, were otherwise completely unexpected. The success of this approach shows that the Darwinian paradigm is capable of explaining not just the adaptations of organisms but also how new kinds of organismal entities come into being.


We thank F. Ayala and J. Avise for organizing the symposium in which this work was presented, John Bonner for kindly providing original data for the social insect portion of Fig. 8.1, and J. J. Boomsma and the anonymous referees for helpful suggestions. This material is based on work supported by National Science Foundation Grant EF0626963.

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