THE SHARPEST critique of Wynne-Edwards came from George C. Williams, in his 1966 book Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. Williams was driven by a distaste for multi-level evolutionary thought that went back to his early years teaching at the University of Chicago in the mid-1950s. The triggering event may have been a lecture by A. E. Emerson, a renowned ecologist and termite specialist. The lecture dealt with what Emerson termed beneficial death, an idea that included August Weismann’s theory that senescence was evolved to cull the old and impaired from populations so that fitter youthful individuals could take their place. My reaction was that if Emerson’s presentation was acceptable biology, I would prefer another calling. (quoted by Borello 2010, p. 107) Likewise, Williams’s tone in a letter to David Lack indicates that he considered the idea of group selection rather ridiculous. You probably had some trouble with the wording of your discussion of Wynne-Edwards. The subject requires great care to avoid the appearance of sarcasm or ridicule. I know that when I got to that part about the epideictic function of the vertical movement of plankton [Wynne-Edwards’s Chapter 16] I suddenly wondered if I had fallen for a really elaborate joke.” (quoted by Borello 2010, p. 111) Williams insisted that adaptations should be attributed to no higher a level of organization than is demanded by the evidence. In explaining adaptation, one should assume the adequacy of the simplest form of natural selection, that of alternative alleles in Mendelian populations, unless the evidence clearly shows that this theory does not suffice. (Williams 1966, pp. 4–5) Conflating the principle of parsimony with the a focus on the small, Williams continued his argument in the following terms: Various levels of adaptive organization, from the subcellular to the biospheric, might conceivably be recognized, but the principle of parsimony demands that we recognize adaptation at the level necessitated by the facts and no higher. It is my position that adaptation need almost never be recognized at any level above that of a pair of parents and associated offspring. (Williams 1966, p. 19)