Examining the costs and motivations of warfare is key to conundrums concerning the relevance of this troubling phenomenon to the evolution of social attachment and cooperation, particularly during adolescence and young adulthood-the developmental time period during which many participants are first recruited for warfare. The study focuses on Samburu, a pastoralist society of approximately 200,000 people occupying northern Kenya's semi-arid and arid lands, asking what role the emotionally sensitized, peer-driven adolescent life stage may have played in the cultural and genetic coevolution of coalitional lethal aggression. Research in small-scale societies provides unparalleled opportunities for sharply defined variables, particularly in age generation societies in which all young men are initiated into "warriorhood." Proposing an epigenetic and component behavior approach, we examine whether raiding activities such as number of raids, killing, and sparing enemy lives associate with DNA methylation in two candidate genes: MAOA, linked to mood and arousal, and NR3C1, linked to stress and immune response. We report statistically significant associations between the epigenetic variables and the combat (exposure) variables of overall raiding activity and reportedly showing mercy to enemies. In contrast, epigenetic variables did not associate with post-traumatic stress disorder (PTSD) symptom scores (a potential outcome measure), and the only combat variable associated with PTSD (but not DNA methylation) was losing one's own livestock in a raid. These findings raise important questions concerning the mechanisms driving warfare's paradoxical mix of violent and altruistic behaviors.
Keywords: Kenya; Warfare; adolescence; cooperation; epigenetics; prosocial emotion.