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transition of the egg from beneath the warmth and safety of the mother to atop the feet and under a fold of feathered abdominal skin of the father requires a bit of coordination on the part of the two. Even this is not sufficient for the genes of this pair to find their way into the gene pool. The conditions of the Antarctic winters are among the least hospitable on earth, with winter temperatures dipping below -60 degrees Celsius and winds reaching 120 mph. To protect themselves from the wind and cold, the male penguins huddle together, spending much of their time sleeping to conserve energy. In this harsh environment, survival of the chicks depends on the shared warmth and protection of the huddle, not the individual. The group as a whole is more likely to survive if each penguin and chick shares in the warmth and protection of the collective structure, which means selective pressures exist to promote cooperation to maintain the integrity of the huddle. More generally, for species born to a period of utter dependency, the genes that find their way into the gene pool are not defined solely or even mostly by likelihood that an organism will reproduce but by the likelihood that the offspring of the parent will live long enough to reproduce. As in the case of the Emperor penguins, one consequence is that selfish genes evolved through individual-level selection processes to promote social preferences and group processes, including reciprocal social behaviors, that can extend beyond kin relationships (e.g., 5). Examples of such selection processes in humans exist, as well (e.g., 6, 7).

The environmental challenges facing Emperor penguins, as daunting as they are, pale by comparison to the complexities facing the human species. Indeed, the social brain hypothesis posits that the social complexities and demands of primate species contributed to the rapid increase in neocortical (i.e., the outer layer of the brain) connectivity and intelligence (8). Warfare among ancestral hunter-gatherers appears to have contributed to group selection for human social behaviors, especially altruistic behaviors (5). As Darwin (1871) noted:

A tribe including many members who, from possessing a high degree of the spirit of patriotism, fidelity, obedience, courage, and sympathy, were always ready to aid one another, and to sacrifice themselves for the common good would be victorious over most other tribes; and this would be natural selection. (p. 166)

Moreover, deducing better ways to find food, avoid perils, and navigate territories has adaptive value for large mammals, but the complexities of these ecological demands are no match for the complexities of social living (especially in hostile between-group social environments), which include: recognizing ingroup and outgroup members; learning by social observation; recognizing the shifting status of friends and foes; anticipating and coordinating efforts between two or more individuals; using language to communicate, reason, teach, and deceive others; orchestrating relationships, ranging from pair bonds and families to friends, bands, and coalitions; navigating complex social hierarchies, social norms and cultural developments; subjugating self-interests to the interests of the pair bond or social group in exchange for the possibility of long term benefits for oneself or one’s group; recruiting support to sanction

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