Birth of the First ESS: George Price, John Maynard Smith, and the Discovery of the Lost "Antlers" Paper

The application of game theory to evolutionary problems is so commonplace today, that few stop to consider how it all began. John Maynard Smith and George R. Price's 1973 Nature article, "The Logic of Animal Conflict," is often referred to as the first description in the literature of the concept of an evolutionary stable strategy (ESS), but what was the "behind the scenes" of the writing of that seminal paper? This article tracks the little known story of the curious American polymath, George Price. As will be shown, it was an earlier paper, the lost "Antlers, Intraspecific Combat, and Altruism," sent by Price to Nature in August 1968 (Unpublished), and refereed by Maynard Smith, which instigated the birth of the first ESS. Recently, the "Antlers" paper has been re-discovered by the author, shedding new light, together with letters and journals from the personal papers of George Price and John Maynard Smith, on their historical paper.