Abstract
Human cooperation (paying a cost to benefit others) is puzzling from a Darwinian perspective, particularly in groups with strangers who cannot repay nor are family members. The beneficial effects of cooperation typically increase nonlinearly with the number of cooperators, e.g., increasing returns when cooperation is low and diminishing returns when cooperation is high. Such nonlinearity can allow cooperation between strangers to persist evolutionarily if a large enough proportion of the population are already cooperators. However, if a lone cooperator faces a conflict between the group's and its own interests (a social dilemma), that raises the question of how cooperation arose in the first place. We use a mathematically tractable evolutionary model to formalise a chronological narrative that has previously only been investigated verbally: given that ancient humans interacted mostly with family members (genetic homophily), cooperation evolved first by kin selection, and then persisted in situations with nonlinear benefits as homophily declined or even if interactions with strangers became the norm. The model also predicts the coexistence of cooperators and defectors observed in the human population (polymorphism), and may explain why cooperators in behavioural experiments prefer to condition their contribution on the contributions of others (conditional cooperation in public goods games).