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We present a simple framework that highlights the most fundamental requirement for the evolution of altruism: assortment between individuals carrying the cooperative genotype and the helping behaviours of others with which these individuals interact. We partition the fitness effects on individuals into those due to self and those due to the 'interaction(More)
Inclusive fitness and reciprocal altruism are widely thought to be distinct explanations for how altruism evolves. Here we show that they rely on the same underlying mechanism. We demonstrate this commonality by applying Hamilton's rule, normally associated with inclusive fitness, to two simple models of reciprocal altruism: one, an iterated prisoner's(More)
Wild et al. argue that the evolution of reduced virulence can be understood from the perspective of inclusive fitness, obviating the need to evoke group selection as a contributing causal factor. Although they acknowledge the mathematical equivalence of the inclusive fitness and multilevel selection approaches, they conclude that reduced virulence can be(More)
If one defines altruism strictly at the population level such that carriers of the altruistic genotype are required to experience, on average, a net fitness cost relative to average population members, then altruism can never evolve. This is simply because a genetically encoded trait can only increase in a population (relative to alternative traits) if the(More)
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(More)
Although the prisoner's dilemma (PD) has been used extensively to study reciprocal altruism, here we show that the n-player prisoner's dilemma (NPD) is also central to two other prominent theories of the evolution of altruism: inclusive fitness and multilevel selection. An NPD model captures the essential factors for the evolution of altruism directly in(More)
reconstructed for dinosaurs are realistic, because other types of curve might fit better, and few data at their lower ends are currently available. A third crucial question is how birds and their immediate dinosaurian relatives became small. Erickson [1], we think, misstates our results [10,11] when he says: 'It was posited that selection favored reduced(More)
The longstanding debate about the importance of group (multilevel) selection suffers from a lack of formal models that describe explicit selection events at multiple levels. Here, we describe a general class of models for two-level evolutionary processes which include birth and death events at both levels. The models incorporate the state-dependent rates at(More)
Hamilton's rule is regarded as a useful tool in the understanding of social evolution, but it relies on restrictive, overly simple assumptions. Here we model more realistic situations, in which the traditional Hamilton's rule generally fails to predict the direction of selection. We offer modifications that allow accurate predictions, but also show that(More)