Altruistic behavior is common throughout the animal kingdom, particularly in species with complex social structures. For example, vampire bats regularly regurgitate blood and donate it to other members of their group who have failed to feed that night, ensuring they do not starve. In numerous bird species, a breeding pair receives help in raising its young from other ‘helper’ birds, who protect the nest from predators and help to feed the fledglings. Vervet monkeys give alarm calls to warn fellow monkeys of the presence of predators, even though in doing so they attract attention to themselves, increasing their personal chance of being attacked. In social insect colonies (ants, wasps, bees and termites), sterile workers devote their whole lives to caring for the queen, constructing and protecting the nest, foraging for food, and tending the larvae. Such behavior is maximally altruistic: sterile workers obviously do not leave any offspring of their own—so have personal fitness of zero—but their actions greatly assist the reproductive efforts of the queen. From a Darwinian viewpoint, the existence of altruism in nature is at first sight puzzling, as Darwin himself realized. Natural selection leads us to expect animals to behave in ways that increase their own chances of survival and reproduction, not those of others. But by behaving altruistically an animal reduces its own fitness, so should be at a selective disadvantage vis-à-vis one which behaves selfishly. To see this, imagine that some members of a group of Vervet monkeys give alarm calls when they see predators, but others do not. Other things being equal, the latter will have an advantage. By selfishly refusing to give an alarm call, a monkey can reduce the chance that it will itself be attacked, while at the same time benefiting from the alarm calls of others. So we should expect natural selection to favour those monkeys that do not give alarm calls over those that do. But this raises an immediate puzzle. How did the alarm-calling behaviour evolve in the first place, and why has it not been eliminated by natural selection? How can the existence of altruism be reconciled with basic Darwinian principles? Dolphins
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