The evolution of altruism through war is highly sensitive to population structure and to civilian and fighter mortality.

The importance of warfare in the evolution of human social behavior remains highly debated. One hypothesis is that intense warfare between groups favored altruism within groups, a hypothesis given some support by computational modeling and, in particular, the work of Choi and Bowles [J.-K. Choi, S. Bowles, Science 318, 636-640 (2007)]. The results of computational models are, however, sensitive to chosen parameter values and a deeper assessment of the plausibility of the parochial altruism hypothesis requires exploring this model in more detail. Here, I use a recently developed method to reexamine Choi and Bowles' model under a much broader range of conditions to those used in the original paper. Although the evolution of altruism is robust to perturbations in most of the default parameters, it is highly sensitive to group size and migration and to the lethality of war. The results show that the degree of genetic differentiation between groups (FST ) produced by Choi and Bowles' original model is much greater than empirical estimates of FST between hunter-gatherer groups. When FST in the model is close to empirically observed values, altruism does not evolve. These results cast doubt on the importance of war in the evolution of human sociality.

Market integration accounts for local variation in generalized altruism in a nationwide lost-letter experiment.

What explains variation in levels of prosocial behavior across communities? And are members of the ingroup and outgroup treated differently? According to evolutionary theories of generalized altruism, market integration should lead to greater levels of prosociality: Market exchange forces people to interact with unknown others, thus creating the conditions for the extension of prosocial behavior beyond close-knit circles to include outgroup members and strangers. Moving away from the evolutionary focus on cross-cultural variation, this article uses the market-integration hypothesis to explain intracultural variation in levels of prosociality in an advanced society. Taking advantage of an ideal setting, this study reports results from a large-scale, nationwide lost-letter experiment in which 5,980 letters were dispersed in a sample of 188 Italian communities. The study confirms the relevance of market integration in accounting for differences in levels of prosociality: In areas where market exchange is dominant, return rates are high. It also casts a light on the relationship between ingroup and outgroup prosociality: Return rates for both Italian and foreign recipients are the same; they vary together; and ingroup returns are highly predictive of outgroup returns at the community level.

Parochial trust and cooperation across 17 societies.

International challenges such as climate change, poverty, and intergroup conflict require countries to cooperate to solve these complex problems. However, the political tide in many countries has shifted inward, with skepticism and reluctance to cooperate with other countries. Thus, cross-societal investigations are needed to test theory about trust and cooperation within and between groups. We conducted an experimental study in 17 countries designed to test several theories that explain why, who, and where people trust and cooperate more with ingroup members, compared with outgroup members. The experiment involved several interactions in the trust game, either as a trustor or trustee. We manipulated partner group membership in the trust game (ingroup, outgroup, or unknown) and if their reputation was at stake during the interaction. In addition to the standard finding that participants trust and cooperate more with ingroup than outgroup members, we obtained findings that reputational concerns play a decisive role for promoting trust and cooperation universally across societies. Furthermore, men discriminated more in favor of their ingroup than women. Individual differences in cooperative preferences, as measured by social value orientation, predicted cooperation with both ingroup and outgroup members. Finally, we did not find support for three theories about the cross-societal conditions that influence the degree of ingroup favoritism observed across societies (e.g., material security, religiosity, and pathogen stress). We discuss the implications for promoting cooperation within and between countries.

Oxytocin reactivity during intergroup conflict in wild chimpanzees.

Intergroup conflict is evident throughout the history of our species, ubiquitous across human societies, and considered crucial for the evolution of humans' large-scale cooperative nature. Like humans, chimpanzee societies exhibit intragroup coordination and coalitionary support during violent intergroup conflicts. In both species, cooperation among group members is essential for individuals to gain access to benefits from engaging in intergroup conflict. Studies suggest that a contributive mechanism regulating in-group cooperation during intergroup conflicts in humans involves the neuropeptide hormone oxytocin, known to influence trust, coordination, and social cognition, although evidence from natural settings is lacking. Here, applying a noninvasive method, we investigate oxytocinergic system involvement during natural intergroup conflicts in wild chimpanzees. We found that chimpanzees of both sexes had significantly higher urinary oxytocin levels immediately before and during intergroup conflict compared with controls. Also, elevated hormone levels were linked with greater cohesion during intergroup conflicts, rather than with the level of potential threat posed by rival groups, intragroup affiliative social interactions, or coordinated behavior alone. Thus, the oxytocinergic system, potentially engendering cohesion and cooperation when facing an out-group threat, may not be uniquely human but rather a mechanism with evolutionary roots shared by our last common ancestor with chimpanzees, likely expediting fitness gains during intergroup conflict.

In-group defense, out-group aggression, and coordination failures in intergroup conflict.

Intergroup conflict persists when and because individuals make costly contributions to their group's fighting capacity, but how groups organize contributions into effective collective action remains poorly understood. Here we distinguish between contributions aimed at subordinating out-groups (out-group aggression) from those aimed at defending the in-group against possible out-group aggression (in-group defense). We conducted two experiments in which three-person aggressor groups confronted three-person defender groups in a multiround contest game (n = 276; 92 aggressor-defender contests). Individuals received an endowment from which they could contribute to their group's fighting capacity. Contributions were always wasted, but when the aggressor group's fighting capacity exceeded that of the defender group, the aggressor group acquired the defender group's remaining resources (otherwise, individuals on both sides were left with the remainders of their endowment). In-group defense appeared stronger and better coordinated than out-group aggression, and defender groups survived roughly 70% of the attacks. This low success rate for aggressor groups mirrored that of group-hunting predators such as wolves and chimpanzees (n = 1,382 cases), hostile takeovers in industry (n = 1,637 cases), and interstate conflicts (n = 2,586). Furthermore, whereas peer punishment increased out-group aggression more than in-group defense without affecting success rates (Exp. 1), sequential (vs. simultaneous) decision-making increased coordination of collective action for out-group aggression, doubling the aggressor's success rate (Exp. 2). The relatively high success rate of in-group defense suggests evolutionary and cultural pressures may have favored capacities for cooperation and coordination when the group goal is to defend, rather than to expand, dominate, and exploit.

Group-level competition influences urinary steroid hormones among wild red-tailed monkeys, indicating energetic costs.

Various theories emphasize that intergroup competition should affect intragroup cooperation and social relationships, especially if the cost of intergroup competition outweighs that of intragroup competition. This cost of intergroup competition may be quantified by changes in physiological status, such as in the steroid hormones cortisol (C) and testosterone (T), which rise or are depressed during periods of energetic stress, respectively. Here we tested for changes in urinary C and T after intergroup encounters (IGEs) among wild red-tailed monkeys (Cercopithecus ascanius), a species that experiences frequent intergroup feeding competition, at the Ngogo station in Kibale National Park, Uganda. We assayed 108 urine samples, of which 36 were collected after IGEs, from 23 individuals in four social groups. Bayesian multilevel models controlling for various confounds revealed that IGEs increased C and decreased T relative to baseline, consistent with an energetic cost to IGEs. The C change was more apparent in samples collected early after IGEs, suggesting an anticipatory increase, whereas the T change was stronger in later samples, suggesting sustained energetic trade-offs. Hormone responses were not affected by the IGE outcome. This cost to intergroup competition, together with little evidence for intragroup competition in redtails and other guenons, establishes an interesting test case for theories emphasizing the effect of intergroup competition on intragroup cooperation.

Threat and parochialism in intergroup relations: lab-in-the-field evidence from rural Georgia.

Competition between groups is widely considered to foster cooperation within groups. Evidence from laboratory experiments hints at the existence of a proximate mechanism by which humans increase their level of cooperation with their ingroup when faced with an external threat. Further work suggests that ingroup cooperation should go along with aggressive behaviour towards the outgroup, although these theories are at odds with others that see high investments in outgroup relations as important means of stabilizing intergroup relations. Surprisingly, few of these arguments have been tested in the field, and existing studies are also limited by the lack of a direct measure of threat perception and aggressive behaviour. This study presents lab-in-the-field results from a rural context where exposure to an ethnic outgroup varies between villages. This context makes it possible to capture levels of threat perception, aggressive behaviour and cooperation without inducing intergroup competition artificially in the laboratory. All concepts are measured behaviourally. In- and outgroup cooperation was measured with a standard public goods game, and a novel experimental protocol was developed that measures perceived threat and aggressive behaviour: the threat game. The results show that levels of perceived threat, ingroup cooperation and aggressive behaviour are higher in regions more strongly exposed to ethnic outsiders. However, exposed regions also show high levels of outgroup cooperation and a concomitant lack of elevated ingroup bias. This pattern is explained by theorizing that communities show parochial altruism when faced with an ethnic outgroup, but balance aggressive behaviour with cooperative offers to diffuse tensions and to keep open channels of mutually beneficial exchange.

Cooperation and conflict: field experiments in Northern Ireland.

The idea that cohesive groups, in which individuals help each other, have a competitive advantage over groups composed of selfish individuals has been widely suggested as an explanation for the evolution of cooperation in humans. Recent theoretical models propose the coevolution of parochial altruism and intergroup conflict, when in-group altruism and out-group hostility contribute to the group's success in these conflicts. However, the few empirical attempts to test this hypothesis do not use natural groups and conflate measures of in-group and unbiased cooperative behaviour. We conducted field experiments based on naturalistic measures of cooperation (school/charity donations and lost letters' returns) with two religious groups with an on-going history of conflict-Catholics and Protestants in Northern Ireland. Conflict was associated with reduced donations to out-group schools and the return of out-group letters, but we found no evidence that it influences in-group cooperation. Rather, socio-economic status was the major determinant of cooperative behaviour. Our study presents a challenge to dominant perspectives on the origins of human cooperation, and has implications for initiatives aiming to promote conflict resolution and social cohesion.

Distinguishing Intergroup and Long-Distance Relationships.

Intergroup and long-distance relationships are both central features of human social life, but because intergroup relationships are emphasized in the literature, long-distance relationships are often overlooked. Here, we make the case that intergroup and long-distance relationships should be studied as distinct, albeit related, features of human sociality. First, we review the functions of both kinds of relationship: while both can be conduits for difficult-to-access resources, intergroup relationships can reduce intergroup conflict whereas long-distance relationships are especially effective at buffering widespread resource shortfalls. Second, to illustrate the importance of distinguishing the two relationship types, we present a case study from rural Bolivia. Combining ethnography and two different experimental techniques, we find that the importance of intergroup relationships-and the salience of group membership itself-varies across populations and across methods. Although ethnography revealed that participants often rely on long-distance relationships for resource access, we were unable to capture participant preferences for these relationships with a forced-choice technique. Taken together, our review and empirical data highlight that (1) intergroup and long-distance relationships can have different functions and can be more or less important in different contexts and (2) validating experimental field data with ethnography is crucial for work on human sociality. We close by outlining future directions for research on long-distance relationships in humans.

Oxytocin has 'tend-and-defend' functionality in group conflict across social vertebrates.

Across vertebrate species, intergroup conflict confronts individuals with a tension between group interests best served by participation in conflict and personal interest best served by not participating. Here, we identify the neurohormone oxytocin as pivotal to the neurobiological regulation of this tension in distinctly different group-living vertebrates, including fishes, birds, rodents, non-human primates and humans. In the context of intergroup conflict, a review of emerging work on pro-sociality suggests that oxytocin and its fish and birds homologues, isotocin and mesotocin, respectively, can elicit participation in group conflict and aggression. This is because it amplifies (i) concern for the interests of genetically related or culturally similar 'in-group' others and (ii) willingness to defend against outside intruders and enemy conspecifics. Across a range of social vertebrates, oxytocin can induce aggressive behaviour to 'tend-and-defend' the in-group during intergroup contests. This article is part of the theme issue 'Intergroup conflict across taxa'.

Environmental stress increases out-group aggression and intergroup conflict in humans.

Peaceful coexistence and trade among human groups can be fragile and intergroup relations frequently transition to violent exchange and conflict. Here we specify how exogenous changes in groups' environment and ensuing carrying-capacity stress can increase individual participation in intergroup conflict, and out-group aggression in particular. In two intergroup contest experiments, individuals could contribute private resources to out-group aggression (versus in-group defense). Environmental unpredictability, induced by making non-invested resources subject to risk of destruction (versus not), created psychological stress and increased participation in and coordination of out-group attacks. Archival analyses of interstate conflicts showed, likewise, that sovereign states engage in revisionist warfare more when their pre-conflict economic and climatic environment were more volatile and unpredictable. Given that participation in conflict is wasteful, environmental unpredictability not only made groups more often victorious but also less wealthy. Macro-level changes in the natural and economic environment can be a root cause of out-group aggression and turn benign intergroup relations violent. This article is part of the theme issue 'Intergroup conflict across taxa'.

The Impact of Positivity and Parochial Altruism on Protective Behaviours during the First COVID-19 Lockdown in Italy.

Implementation of COVID-19 protective behaviours, such as social distancing or frequent hand washing during the lockdown, was critical to prevent the spread of the COVID-19 pandemic. In this cross-sectional study, we examined the effect of positivity and parochial altruism on implementing COVID-19 health-protective behaviours during the Italian lockdown. A sample of 460 participants completed an online questionnaire that included demographic measures, Positivity Scale and COVID-19 measures of health-protective behaviours. To measure parochial altruism, we used a hypothetical dictator game played with others who could vary in their social distance from the participants. Results showed that participants in the hypothetical game gave more money to parents and siblings than to best friends, cousins, neighbours, and strangers. Furthermore, both positivity and parochial altruism (more altruism toward close vs. distant people) were positively associated with implementing hygiene behaviours but not with social distancing. Finally, mediation analysis showed that increases in parochial altruism mediated the effect of positivity on hygiene behaviour. These findings extend knowledge about the factors beyond the implementation of COVID-19 health-protective behaviours during a lockdown situation.

Political ideology, cooperation and national parochialism across 42 nations.

Political ideology has been hypothesized to be associated with cooperation and national parochialism (i.e. greater cooperation with members of one's nation), with liberals thought to have more cooperation with strangers and less national parochialism, compared to conservatives. However, previous findings are limited to few-and predominantly western-nations. Here, we present a large-scale cross-societal experiment that can test hypotheses on the relation between political ideology, cooperation and national parochialism around the globe. To do so, we recruited 18 411 participants from 42 nations. Participants made decisions in a prisoner's dilemma game, and we manipulated the nationality of their interaction partner (national ingroup member, national outgroup member or unidentified stranger). We found that liberals, compared to conservatives, displayed slightly greater cooperation, trust in others and greater identification with the world as a whole. Conservatives, however, identified more strongly with their own nation and displayed slightly greater national parochialism in cooperation. Importantly, the association between political ideology and behaviour was significant in nations characterized by higher wealth, stronger rule of law and better government effectiveness. We discuss the implications of these findings for understanding the association between political ideology and cooperation. This article is part of the theme issue 'The political brain: neurocognitive and computational mechanisms'.

Ethnic Markers without Ethnic Conflict : Why do Interdependent Masikoro, Mikea, and Vezo of Madagascar Signal their Ethnic Differences?

People often signal their membership in groups through their clothes, hairstyle, posture, and dialect. Most existing evolutionary models argue that markers label group members so individuals can preferentially interact with those in their group. Here we ask why people mark ethnic differences when interethnic interaction is routine, necessary, and peaceful. We asked research participants from three ethnic groups in southwestern Madagascar to sort photos of unfamiliar people by ethnicity, and by with whom they would prefer or not prefer to cooperate, in a wage labor vignette. Results indicate that southwestern Malagasy reliably send and detect ethnic signals; they signal less in the marketplace, a primary site of interethnic coordination and cooperation; and they do not prefer co-ethnics as cooperation partners in novel circumstances. Results from a cultural knowledge survey and calculations of cultural FST suggest that these ethnic groups have relatively little cultural differentiation. We concur with Moya and Boyd (Human Nature 26:1-27, 2015) that ethnicity is unlikely to be a singular social phenomenon. The current functions of ethnic divisions and marking may be different from those at the moment of ethnogenesis. Group identities may persist without group conflict or differentiation.

RéSUMé: Les gens montrent souvent leur appartenance à un groupe à travers leurs modes vestimentaires, leur style de coiffure, leur posture et surtout leur dialecte. La plupart des modèles évolutifs existants soutiennent que les marqueurs caractérisent les membres du groupe afin que les individus puissent interagir de manière préférentielle avec les membres de leur groupe. Nous nous demandons ici, pourquoi les gens marquent les différences ethniques lorsque l'interaction interethnique est routinière, nécessaire et pacifique. Nous avons alors demandé à des participants issus de trois groupes ethniques du Sud-Ouest de Madagascar de trier des photos de personnes inconnues en fonction de leur appartenance ethnique, et en fonction des personnes avec lesquelles ils préféreraient ou non coopérer, dans une vignette hypothétique. Les résultats recueilli indiquent clairement que les Malgaches du Sud-Ouest émettent et détectent de manière fiable les indicateurs ethniques; ils émettent moins de signaux indicatifs sur la place du marché, dans un site primaire de coordination et de coopération interethnique; et ils ne préfèrent pas les co-ethnies comme partenaires de coopération dans des circonstances nouvelles. Basé sur les résultats obtenus d'une enquête réalisée sur les connaissances culturelles et les calculs du FST culturel suggèrent que ces groupes ethniques présentent une différenciation culturelle relativement faible. Nous partageons l'opinion de Moya et Boyd (Human Nature 26:1­27, 2015) pour dire qu'il est peu probable que l'ethnicité soit un phénomène social singulier. Les fonctions actuelles des divisions et du marquage ethniques peuvent être différentes de celles du moment de l'ethnogenèse. Les identités de groupe peuvent persister sans qu'il y ait conflit ou différenciation de groupe.

The demography of human warfare can drive sex differences in altruism.

Recent years have seen great interest in the suggestion that between-group aggression and within-group altruism have coevolved. However, these efforts have neglected the possibility that warfare - via its impact on demography - might influence human social behaviours more widely, not just those directly connected to success in war. Moreover, the potential for sex differences in the demography of warfare to translate into sex differences in social behaviour more generally has remained unexplored. Here, we develop a kin-selection model of altruism performed by men and women for the benefit of their groupmates in a population experiencing intergroup conflict. We find that warfare can promote altruistic, helping behaviours as the additional reproductive opportunities winners obtain in defeated groups decrease harmful competition between kin. Furthermore, we find that sex can be a crucial modulator of altruism, with there being a tendency for the sex that competes more intensely with relatives to behave more altruistically and for the sex that competes more intensely with non-relatives in defeated groups to receive more altruism. In addition, there is also a tendency for the less-dispersing sex to both give and receive more altruism. We discuss implications for our understanding of observed sex differences in cooperation in human societies.

Oxytocin modulates third-party sanctioning of selfish and generous behavior within and between groups.

Human groups function because members trust each other and reciprocate cooperative contributions, and reward others' cooperation and punish their non-cooperation. Here we examined the possibility that such third-party punishment and reward of others' trust and reciprocation is modulated by oxytocin, a neuropeptide generally involved in social bonding and in-group (but not out-group) serving behavior. Healthy males and females (N=100) self-administered a placebo or 24 IU of oxytocin in a randomized, double-blind, between-subjects design. Participants were asked to indicate (incentivized, costly) their level of reward or punishment for in-group (outgroup) investors donating generously or fairly to in-group (outgroup) trustees, who back-transferred generously, fairly or selfishly. Punishment (reward) was higher for selfish (generous) investments and back-transfers when (i) investors were in-group rather than outgroup, and (ii) trustees were in-group rather than outgroup, especially when (iii) participants received oxytocin rather than placebo. It follows, first, that oxytocin leads individuals to ignore out-groups as long as out-group behavior is not relevant to the in-group and, second, that oxytocin contributes to creating and enforcing in-group norms of cooperation and trust.

Deciphering the modulatory role of oxytocin in human altruism.

Unlike any other species, humans frequently engage in altruistic behaviors by which they increase another individual's welfare even if this implies personal costs. The psychological motives underlying altruistic behaviors remain diverse, ranging from the ability to reciprocate trust and cooperation to bonding and empathizing with family members or even genetically unrelated others. This article explores the neuroendocrine architecture of altruism by emphasizing the crucial role of the evolutionarily highly conserved peptide hormone oxytocin as a modulator of cooperative behaviors including empathy-driven altruism. However, accumulating evidence suggests that oxytocin does not invariably facilitate cooperation but also produces protective or even defensive-aggressive responses in specific social contexts. In addition, we highlight the relevance of message frames as critical determinants of whether the peptide promotes altruism toward prosocial ends.

An experiment on individual 'parochial altruism' revealing no connection between individual 'altruism' and individual 'parochialism'.

Is parochial altruism an attribute of individual behavior? This is the question we address with an experiment. We examine whether the individual pro-sociality that is revealed in the public goods and trust games when interacting with fellow group members helps predict individual parochialism, as measured by the in-group bias (i.e., the difference in these games in pro-sociality when interacting with own group members as compared with members of another group). We find that it is not. An examination of the Big-5 personality predictors of each behavior reinforces this result: they are different. In short, knowing how pro-social individuals are with respect to fellow group members does not help predict their parochialism.

Dying the right-way? Interest in and perceived persuasiveness of parochial extremist propaganda increases after mortality salience.

Research on parochial altruism demonstrated that hostility toward out-groups (parochialism) represents the dark side of the willingness to benefit one's in-group even at own costs (altruism). Parochial aggression thereby emerged mainly under conditions of threat. Extremist propaganda videos, for instance by right-wing extremists, try to capitalize on parochial altruistic mechanism by telling recipients sharing their national identity that this nation is under threat wherefore they for have to join the extremist's cause to prevent the extinction of their nation. Most of the time, propaganda videos are rated as uninteresting and non-persuasive by the target audience. Yet, evolutionary media psychology posits that the interest in and effectiveness of media increases when evolutionarily relevant problems are addressed. Consequently, interest in parochial altruistic right-wing extremist messages should increase under conditions of threat. The current study tested this assumption by randomly assigning German non-Muslims (N = 109) to either an existential threat (here: mortality salience) or a control condition and asking them to evaluate extremist propaganda that addressed them as either in-group members (right-wing extremists) or as out-group members (Islamic extremists). In support of the hypotheses, subjects under conditions of threat reported a higher interest in the right-wing extremist propaganda and perceived it as more persuasive. We discuss the results concerning the implications for evolutionary media psychology and the transmission of parochial altruism in propaganda videos.

In intergroup conflict, self-sacrifice is stronger among pro-social individuals, and parochial altruism emerges especially among cognitively taxed individuals.

Parochial altruism is decomposed in a tendency to benefit the in-group along with a tendency to ignore, derogate, and harm rivaling out-groups. Building off recent work suggesting that decisions to cooperate can be relatively fast and intuitive, we examine parochial altruism in intergroup conflict when cognitive deliberation is rendered difficult or not. Predictions were tested in an experiment using an incentivized Intergroup Prisoner's Dilemma-Maximizing Differences Game with 95 subjects classified as either pro-social or pro-self being randomly allocated to high vs. low impulse-control conditions. Results showed, first of all, that self-sacrificial decisions to contribute were made faster than decisions not to contribute, and that faster decision time associated with more positive expectations of in-group members. Second, we observed that lowering impulse control with a difficult rather than easy Stroop Task increased the amount contributed to a pool that benefited in-group members while harming out-group members; thus reducing deliberation increased parochial altruism. Finally, results replicated earlier work showing that especially pro-social (vs. pro-self) individuals contributed more to the in-group and did not lower their contributions to the between-group pool that benefitted their in-group and, simultaneously, hurt the out-group. This pattern emerged independent of their impulse control. Thus, (in-group bounded) cooperation is more prominent among individuals with strong rather than weak other-regarding preferences. Moreover, the intuitive tendency to cooperate may have evolved in the context of intergroup conflict and therefore is sharp-edged-in-group bounded and including willingness to aggress out-groups.