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Despite the global spread of intensive agriculture, many populations retained foraging or mixed subsistence strategies until well into the twentieth century. Understanding why has been a longstanding puzzle. One explanation, called the marginal habitat hypothesis, is that foraging persisted because foragers tended to live in marginal habitats generally not suited to agriculture. However, recent empirical studies have not supported this view. The alternative but untested oasis hypothesis of agricultural intensification claims that intensive agriculture developed in areas with low biodiversity and a reliable water source not reliant on local rainfall. We test both the marginal habitat and oasis hypotheses using a cross-cultural sample drawn from the 'Ethnographic atlas' (Murdock 1967 Ethnology 6 , 109-236). Our analyses provide support for both hypotheses. We found that intensive agriculture was unlikely in areas with high rainfall. Further, high biodiversity, including pathogens associated with high rainfall, appears to have limited the development of intensive agriculture. Our analyses of African societies show that tsetse flies, elephants and malaria are negatively associated with intensive agriculture, but only the effect of tsetse flies reached significance. Our results suggest that in certain ecologies intensive agriculture may be difficult or impossible to develop but that generally lower rainfall and biodiversity is favourable for its emergence. This article is part of the theme issue 'Evolutionary ecology of inequality'.

Intergroup violence is challenging to understand: why do individuals cooperate to harm members of other groups when they themselves may be killed or injured? Despite progress in understanding the evolutionary and proximate mechanisms that underlie violence, we still have little insight into the processes that lead to the emergence of coalitionary aggression. We argue that an overlooked component is the presence of individuals who have a crucial role in initiating violence. In instigating intergroup violence, these key individuals may expect to face lower costs, receive greater benefits, or garner benefits that have a greater value to them than others. Alternatively, key individuals may be motivated by individual traits such as increased boldness, propensity for aggression or exploratory behaviour. Key individuals catalyse the emergence of coalitionary violence through one of several processes including altering the costs and benefits that accrue to others, paying a greater share of the startup costs, signalling privileged knowledge, or providing coordination, among other factors. Here we integrate diverse lines of empirical research from humans and non-human animals demonstrating that inter-individual variation is an important factor in the emergence of intergroup violence. Focusing on the role of key individuals provides new insights into how and why violence emerges. This article is part of the theme issue 'Intergroup conflict across taxa'.

Humans often favour ingroup members over others, a bias that drives discrimination and intergroup conflicts. Hostile relations between groups and homogeneity within groups may affect such ingroup bias. In an experiment with members of three natural groups in Ethiopia, we vary intergroup relations (neutral versus enmity) and exploit the natural variation in the homogeneity of groups (homogeneous versus heterogeneous) to identify their effect on in- and outgroup concerns. We find that ingroup bias largely manifests as positive concern for ingroup members combined with no concern for outgroup members. Enmity has no effect on ingroup bias, whereas ingroup concern is amplified in homogeneous groups. Group homogeneity, thus, is the primary driver of concerns for others in our study's context. Our results are relevant to understanding the consequences of exclusionary group identities. This article is part of the theme issue 'Intergroup conflict across taxa'.

Models of collective action infrequently account for differences across individuals beyond a limited set of strategies, ignoring variation in endowment (e.g. physical condition, wealth, knowledge, personality, support), individual costs of effort, or expected gains from cooperation. However, behavioural research indicates these inter-individual differences can have significant effects on the dynamics of collective action. The papers contributed to this theme issue evaluate how individual differences affect the propensity to cooperate, and how they can catalyse others' likelihood of cooperation (e.g. via leadership). Many of the papers emphasize the relationship between individual decisions and socio-ecological context, particularly the effect of group size. All together, the papers in this theme issue provide a more complete picture of collective action, by embracing the reality of inter-individual variation and its multiple roles in the success or failure of collective action., (© 2015 The Author(s).)

Observation of leadership in small-scale societies offers unique insights into the evolution of human collective action and the origins of sociopolitical complexity. Using behavioural data from the Tsimane forager-horticulturalists of Bolivia and Nyangatom nomadic pastoralists of Ethiopia, we evaluate the traits of leaders and the contexts in which leadership becomes more institutional. We find that leaders tend to have more capital, in the form of age-related knowledge, body size or social connections. These attributes can reduce the costs leaders incur and increase the efficacy of leadership. Leadership becomes more institutional in domains of collective action, such as resolution of intragroup conflict, where collective action failure threatens group integrity. Together these data support the hypothesis that leadership is an important means by which collective action problems are overcome in small-scale societies., (© 2015 The Author(s).)

Life abounds with examples of conspecifics actively cooperating to a common end, despite conflicts of interest being expected concerning how much each individual should contribute. Mathematical models typically find that such conflict can be resolved by partial-response strategies, leading investors to contribute relatively equitably. Using a case study approach, we show that such model expectations can be contradicted in at least four disparate contexts: (i) bi-parental care; (ii) cooperative breeding; (iii) cooperative hunting; and (iv) human cooperation. We highlight that: (a) marked variation in contributions is commonplace; and (b) individuals can often respond positively rather than negatively to the contributions of others. Existing models have surprisingly limited power in explaining these phenomena. Here, we propose that, although among-individual variation in cooperative contributions will be influenced by differential costs and benefits, there is likely to be a strong genetic or epigenetic component. We then suggest that selection can maintain high investors (key individuals) when their contributions promote support by increasing the benefits and/or reducing the costs for others. Our intentions are to raise awareness in--and provide testable hypotheses of--two of the most poorly understood, yet integral, questions regarding cooperative ventures: why do individuals vary in their contributions and when does cooperation beget cooperation?, (© 2015 The Author(s).)