Hunter-Gatherers Maintain Assortativity in Cooperation despite High Levels of Residential Change and Mixing.

Widespread cooperation is a defining feature of human societies from hunter-gatherer bands to nation states [1, 2], but explaining its evolution remains a challenge. Although positive assortment of cooperators is recognized as a basic requirement for the evolution of cooperation, the mechanisms governing assortment are debated. Moreover, the social structure of modern hunter-gatherers, characterized by high mobility, residential mixing, and low genetic relatedness [3], undermines assortment and adds to the puzzle of how cooperation evolved. Here, we analyze four years of data (2010, 2013, 2014, 2016) tracking residence and levels of cooperation elicited from a public goods game in Hadza hunter-gatherers of Tanzania. Data were collected from 56 camps, comprising 383 unique individuals, 137 of whom we have data for two or more years. Despite significant residential mixing, we observe a robust pattern of assortment that is necessary for cooperation to evolve; in every year, Hadza camps exhibit high between-camp and low within-camp variation in cooperation. We find little evidence that cooperative behavior within individuals is stable over time or that similarity in cooperation between dyads predicts their future cohabitation. Both sets of findings are inconsistent with models that assume stable cooperative and selfish types, including partner choice models. Consistent with social norms, culture, and reciprocity theories, the strongest predictor of an individual's level of cooperation is the mean cooperation of their current campmates. These findings underscore the adaptive nature of human cooperation-particularly its responsiveness to social contexts-as a feature that is important in generating the assortment necessary for cooperation to evolve.