Obstacles and catalysts of cooperation in humans, bonobos, and chimpanzees: behavioural reaction norms can help explain variation in sex roles, inequality, war and peace

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Our closest living relatives, bonobos and chimpanzees, along with small-scale human societies figure prominently in debates about human nature. Here we emphasize and explain behavioural variation within and among these three species. In the logic of behavioural ecology, individuals have been selected to adjust their behaviour along evolved reaction norms that maximize fitness given current socio-ecological conditions. We discuss variation in three behavioural contexts: relationships between the sexes, hierarchy and inequality, and intergroup interactions. In each context, behavioural variation can be related to two broad socio-ecological conditions: (i) the defensibility of contested resources, and (ii) differences in bargaining power. When defensibility of resources and differences in bargaining power are great, interactions are rife with conflict; when they are minimal, interactions are more harmonious. These socio-ecological conditions therefore constitute key catalysts and obstacles of cooperation. We conclude that human nature should be seen as consisting of evolved reaction norms.

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Figures
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    Behavioural reaction norms express how individuals respond to variation in the environment (x-axis) by adaptive changes in behavioural strategy (y-axis) (Dingemanse et al., 2010). Individuals or species can differ in their average behaviour (elevation of the reaction norm) and/or their behavioural plasticity (slope) (Jaeggi et al., 2010a); here, two hypothetical individuals or species are shown that differ in both (see Dingemanse et al., 2010 for discussion of this and other examples). Ultimately, species differences in elevation or slope should correspond to differences in the fitness benefits of the behavioural strategy and the expected range of environmental variation as calibrated by ancestral environments. Proximately, shifts along these behavioural reaction norms could, e.g., be mediated by neuroendocrine mechanisms, through changes in baseline hormone levels (elevation) or acute reactivity (slope) (Trumble et al., 2015). For simplicity we here adopt linear reaction norms though other shapes are plausible. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1568539x.

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    Proposed variation in mating patterns and sexual selection among chimpanzees and bonobos as a function of food availability. Primary effects of higher food availability (less seasonality, larger patches) are to increase female sociality by alleviating feeding competition and thereby increasing female power through coalitions, as well as to increase female sexual activity (by alleviating the costs of mating and reproduction) and thereby levelling the operational sex ratio. Secondary effects then are increased female mate choice (blue line) and decreased male mating competition (red line). Shaded areas at the top indicate the suggested chimpanzee and bonobo ranges, with Gombe and Taï as extreme examples. For simplicity, the two species are represented as having identical, linear reaction norms. See Section 2.2 for details and Table 1 for data on female sociality and sex ratios. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1568539x.

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    Variation in human marriage, mating and parenting as arising from variation in male bargaining power. As men gain bargaining power (through increased control over the means of production, greater mate quality, a female-biased sex ratio, or more social support), their contribution to existing partnerships decreases (blue line). With lower male parenting effort, monogamous marriage becomes less stable, and male pursuit of additional matings increase, resulting in higher levels of polygyny at the individual and population level (red line). See Section 2.3 for examples and details. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1568539x.

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    Across species, the extent to which resources are defensible should positively associate with resultant inequality as dominants are able to hoard larger shares. However, the elevation of this reaction norm can decrease if subordinates maintain leverage due to a need for coalitions in resource defence, cooperative production, outside options, or reduced differences in fighting ability. The slope might decrease as depicted here if the importance of collective action, and hence subordinate leverage, in procuring and protecting resources increases with defensibility as might be the case for complex foragers and chimpanzees. Placement of example populations is tentative, see Section 3 for details. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1568539x.

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    Intergroup correlation in production (resource abundance) changes the gains to raiding or trading. If the correlation is low, one group will have plenty when others have little and vice versa, creating the opportunity to benefit from reciprocal land use and trade, e.g., among most foragers and perhaps bonobos and Taï chimpanzees where females may visit neighbouring groups for extended periods of time. When neighbours experience good and bad times simultaneously, this increases the gains to territoriality and warfare, e.g., among complex foragers and agriculturalists. The benefits from both kinds of intergroup interactions are reduced when the risk of shortfalls is lower (dashed vs solid lines). After Kelly (2013), his Figures 6–8. See Section 4 for details and examples. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1568539x.

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