Reciprocity creates a stake in one's partner, or why you should cooperate even when anonymous

Why do we care so much for friends, even making sacrifices for them they cannot repay or never know about? When organisms engage in reciprocity, they have a stake in their partner's survival and wellbeing so the reciprocal relationship can persist. This stake (aka fitness interdependence) makes organisms willing to help beyond the existing reciprocal arrangement (e.g. anonymously). I demonstrate this with two mathematical models in which organisms play a prisoner's dilemma, and where helping keeps their partner alive and well. Both models shows that reciprocity creates a stake in partners' welfare: those who help a cooperative partner--even when anonymous--do better than those who do not, because they keep that cooperative partner in good enough condition to continue the reciprocal relationship. 'Machiavellian' cooperators, who defect when anonymous, do worse because their partners become incapacitated. This work highlights the fact that reciprocity and stake are not separate evolutionary processes, but are inherently linked.