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Behavioral Ecology Vol. 12 No. 3: 261
© 2001 International Society for Behavioral Ecology


Hamilton Symposium

Introduction to William D. Hamilton Symposium

Stephen T. Emlen

Past President, International Society for Behavioral Ecology J. G. Schurman Professor of Behavioral Ecology, Cornell University

On 7 March 2000, William Donald Hamilton died in Oxford, England, from malaria contracted during a field expedition to the Congo. With his death, the field of evolutionary biology arguably lost its greatest practitioner since Charles Darwin. W. D. Hamilton made major contributions to many areas of evolutionary biology, but perhaps his greatest impact was on the discipline of behavioral ecology. The topics that he focused his energies upon include kin selection, reciprocity, sex-ratio selection, the evolution of senescence, alternative reproductive strategies, selfish gains of group living, haplodiploidy and the evolution of eusociality, parasite—host arms races, and the role of parasites in both mate choice and the evolution of sex. His contributions formed a foundation for the new field of sociobiology, and his theories provide the bases for a good deal of the theoretical and empirical work currently being done in behavioral ecology.

Given the intellectual contributions of the man and the magnitude of the void caused by his loss, the International Society for Behavioral Ecology decided to honor the life of William Hamilton in two tangible ways. The first was to hold a symposium in his honor at the Eighth International Behavioral Ecology Congress in Zurich, Switzerland, on 10 August 2000. The proceedings of the symposium are published in the pages that follow. The second is to sponsor a William D. Hamilton Lectureship to be presented at each future ISBE Congress. The society's executive council will biennially select a distinguished evolutionary behaviorist to present this lecture.

William Hamilton was, first and foremost, a passionate natural historian. He traveled throughout the world, but especially loved its tropical regions. He repeatedly stated that he obtained many of his research ideas from simple observations made in nature. Yet Hamilton was no ordinary natural historian. He had the knack of selecting research topics that had broad evolutionary implications. He sought out unanswered questions—evolutionary paradoxes—that were not readily explicable by current natural selection theory. As Richard D. Alexander so aptly expressed it, "Bill's originality of mind often turned the barely articulated ideas of distinguished predecessors—ideas overlooked or neglected by all the rest of us—into magnificent theoretical edifices affecting our view of all life" (Natural History, vol. 6, 2000, pp. 44-46).

Hamilton may have taken his ideas from nature, but he developed his hypotheses in the form of rigorous mathematical models. In an interview with Frans Roes, he said: "Often I use mathematics because I need to straighten out my own ideas. I have a somewhat illogical brain, and unless I put it through the mill of mathematics, I can continue to believe in the impossible for a long time" (International Society of Human Ethology Newsletter, vol. 12, 1997, pp. 3-7). The math helped to make his assumptions clear and his predictions more precise. This, in turn, enabled others not only to understand his innovative ideas but also to design better empirical tests of his theories.

For our symposium, we selected three of the theoretical edifices that owe much to William Hamilton: kin selection and social evolution, sexual selection and parasites, and parasites and the evolution of sex. We asked David Queller, Manfred Milinski, and Paul Schmid-Hempel to summarize and comment on Hamilton's contributions to each area. We trust that readers will find the symposium papers to be valuable personalized synopses, not only of Hamilton's contributions, but also of current unresolved issues in each area.

It is too soon to measure the full magnitude of William Hamilton's impact on evolutionary biology, but it is safe to say that it will be immense. We can state with certainty that his ideas on inclusive fitness and reciprocity have revolutionized the ways in which we view social conflict and cooperation. The jury currently is more mixed on the relative importance of his thoughts concerning parasite—host arms races as a primary cause for the evolution of sexual reproduction. But there can be little doubt that succeeding generations of evolutionary biologists in general, and behavioral ecologists in particular, will continue to find fertile ground in the innovative ideas first suggested or formalized by Hamilton. He will be sorely missed. We hope that the symposium papers that follow will help readers to put some of Hamilton's contributions into perspective.


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Proc. Natl. Acad. Sci. USAHome page
W. J. Alonso and C. Schuck-Paim
Sex-ratio conflicts, kin selection, and the evolution of altruism
PNAS, May 14, 2002; 99(10): 6843 - 6847.
[Abstract] [Full Text] [PDF]


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