摘要:Moral rules allow humans to cooperate by indirect reciprocity. Yet, it is not clear which moral rules best implement indirect reciprocity and are favoured by natural selection. Previous studies either considered only public assessment, where individuals are deemed good or bad by all others, or compared a subset of possible strategies. Here we fill this gap by identifying which rules are evolutionary stable strategies (ESS) among all possible moral rules while considering private assessment. We develop an analytical model describing the frequency of long-term cooperation, determining when a strategy can be invaded by another. We show that there are numerous ESSs in absence of errors, which however cease to exist when errors are present. We identify the underlying properties of cooperative ESSs. Overall, this paper provides a first exhaustive evolutionary invasion analysis of moral rules considering private assessment. Moreover, this model is extendable to incorporate higher-order rules and other processes.