Eusocial Insects as a Model for Understanding Altruism, Cooperation, and Levels of Selection
If the logic of natural selection is applied strictly at the level of individual production of offspring, sterile workers in insect societies are enigmatic. How can natural selection ever produce individuals that refrain from reproduction, and how are traits of such individuals that never produce offspring scrutinized and changed through natural selection? The solution to both questions is found in the family structures of insect societies. That is, the sterile helper individuals are evolutionary altruists that give up their own reproduction and instead are helping their kin reproduce and proliferate shared genes in the offspring of the fertile queen. Selection in such cases is not just a matter of individual’s direct reproduction, and instead of own offspring, the currency of the evolutionary success of sterile individuals is inclusive fitness. The concept of inclusive fitness and the process of kin selection are key to understanding the magnificent cooperation we see in insect societies, and reciprocally, insect societies are key case studies of inclusive fitness logic. In extreme cases, such as the highly advanced and sophisticated societies of ants, honeybees, and termites, the division of labor and interdependence of colony members is so complete, that it is justified to talk about a new level of evolutionary individuality. Such increases in the hierarchical complexity of life are called major transitions in evolution. We see adaptations of the colony, rather than individuals, in, e.g., their communication and group behaviors. The division of labor between morphologically differentiated queens and workers is analogous to germline-soma separation of a multicellular organism, justifying the term superorganism for the extreme cases of social lifestyle. Alongside these extreme cases, there is enormous diversity in the social lifestyles across social insect taxa, which provides a window into the balance of cooperation and conflict, and individual reproduction and helping others, in social evolution. Over the last decades, social insect research has been an area where the theoretical and empirical understanding have been developed hand in hand, together with examples of wonderful natural history, and has tremendously improved our understanding of evolution.