Abstract: Although positive reciprocity (reciprocal altruism) has been a focus of interest in evolutionary biology, negative reciprocity (retaliatory infliction of fitness reduction) has been largely ignored. In social animals, retaliatory aggression is common, individuals often punish other group members that infringe their interests, and punishment can cause subordinates to desist from behaviour likely to reduce the fitness of dominant animals. Punishing strategies are used to establish and maintain dominance relationships, to discourage parasites and cheats, to discipline offspring or prospective sexual partners and to maintain cooperative behaviour.