Journal of Theoretical Biology, 228:303-313, 2004.
Although the conditions under which altruistic behaviors evolve continue to be vigorously debated, there is general agreement that altruistic traits involving an absolute cost to altruists (strong altruism) cannot evolve when populations are structured with randomly formed groups. This conclusion implies that the evolution of such traits depends upon special environmental conditions or additional organismic capabilities that enable altruists to interact with each other more than would be expected with random grouping. Here we show, using both analytic and simulation results, that the positive assortment necessary for strong altruism to evolve does not require these additional mechanisms, but merely that randomly formed groups exist for more than one generation. Conditions favoring the selection of altruists, which are absent when random groups initially form, can naturally arise even after a single generation within groups—and even as the proportion of altruists simultaneously decreases. The gains made by altruists in a second generation within groups can more than compensate for the losses suffered in the first and in this way altruism can ratchet up to high levels. This is true even if altruism is initially rare, migration between groups allowed, homogeneous altruist groups prohibited, population growth restricted, or kin selection precluded. Until now random group formation models have neglected the significance of multigenerational groups—even though such groups are a central feature of classic "haystack" models of the evolution of altruism. We also explore the important role that stochasticity (effectively absent in the original infinite models) plays in the evolution of altruism. The fact that strong altruism can increase when groups are periodically and randomly formed suggests that altruism may evolve more readily and in simpler organisms than is generally appreciated.