Intergroup Contests Research Articles

Governance and competition

When groups compete against each other in contests or tournaments they typically differ with regard to the way they are organized and how decisions within groups are determined. In this paper, I experimentally investigate the impact of a group’s organizational structure on inter-group contests. My results show that letting group members decide autonomously leads to significantly lower levels of competition compared to when groups are organized democratically or autocratically. Contrary to my theoretical predictions, I observe no differences between democratically and autocratically organized groups. One reason for this finding is that many individuals in the role of autocratic decision-makers do not use their power to fully exploit their subordinates. Despite this, I find that when giving group members the choice, most individuals prefer the democratic regime, which guarantees them participation in the decision-making process and protects them from exploitation.

Fighting force and experience combine to determine contest success in a warlike mammal

Conflicts between social groups or "intergroup contests" are proposed to play a major role in the evolution of cooperation and social organization in humans and some nonhuman animal societies. In humans, success in warfare and other collective conflicts depends on both fighting group size and the presence and actions of key individuals, such as leaders or talismanic warriors. Understanding the determinants of intergroup contest success in other warlike animals may help to reveal the role of these contests in social evolution. Using 19 y of data on intergroup encounters in a particularly violent social mammal, the banded mongoose (Mungos mungo), we show that two factors, the number of adult males and the age of the oldest male (the "senior" male), have the strongest impacts on the probability of group victory. The advantage conferred by senior males appears to stem from their fighting experience. However, the galvanizing effect of senior males declines as they grow old until, at very advanced ages, senior males become a liability rather than an asset and can be evicted. As in human conflict, strength in numbers and the experience of key individuals combine to determine intergroup contest success in this animal society. We discuss how selection arising from intergroup contests may explain a suite of features of individual life history and social organization, including male eviction, sex-assortative alloparental care, and adult sex ratio.

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’.

Brood as booty: the effect of colony size and resource value in social insect contests

AbstractAnimals engage in contests for access to resources like food, mates, and space. Intergroup contests between groups of organisms have received little attention, and it remains unresolved what information groups might use collectively to make contest decisions. We staged whole-colony contests using ant colonies (Temnothorax rugatulus), which perceive conspecific colonies as both a threat and resource from which to steal brood. We recorded individual behaviors and used demographic characteristics as proxies for resource value (number of brood items) and fighting ability (number of workers). We found that ants altered their fighting effort depending on the relative number of workers of their opponent. Although the proximate mechanism for this ability remains uncertain, we found that colonies increased fighting when their opponent had relatively more brood, but not if opposing colonies had relatively many more workers. This suggests that ant colonies can use information about opposing colonies that shapes contest strategies. Further, the behavior of opposing colonies was strongly correlated with each other despite colony size differences ranging from 4% to 51%, consistent with the hypothesis that colonies can use opponent information. The behavior of a distributed, collective system of many individuals, like a eusocial insect colony, thus fits several predictions of contest models designed for individuals if we consider the gain and loss of worker ants analogous to energetic costs accrued during typical dyadic contests.

Sex (similarities and) differences in friendship jealousy

Friendships provide material benefits, bolster health, and may help solve adaptive challenges. However, a recurrent obstacle to sustaining those friendships—and thus enjoying many friendship-mediated fitness benefits—is interference from other people. Friendship jealousy may be well-designed for helping both men and women meet the recurrent, adaptive challenge of retaining friends in the face of such third-party interference. Although we thus expect several sex similarities in the general cognitive architecture of friendship jealousy (e.g., it is attuned to friend value), there are also sex differences in friendship structures and historical functions, which might influence the inputs of friendship jealousy (e.g., the value of any one friendship). If so, we should also expect some sex differences in friendship jealousy. Findings from a reanalysis of previously-published data and a new experiment, including both U.S. student and adult community participants (N = 993), provide initial support for three predicted sex differences: women (versus men) report greater friendship jealousy at the prospective loss of best friends to others, men (versus women) report greater friendship jealousy at the prospective loss of acquaintances to others, and men's (but not women's) friendship jealousy is enhanced in the context of intergroup contests.

Not all group members are created equal: heterogeneous abilities in inter-group contests

Competition between groups is ubiquitous in social and economic life, and typically occurs between groups that are not created equal. Here we experimentally investigate the implications of this general observation on the unfolding of symmetric and asymmetric competition between groups that are either homogeneous or heterogeneous in the ability of their members to contribute to the success of the group. Our main finding is that relative to the benchmark case in which two homogeneous compete against each other, heterogeneity within groups per se has no discernable effect on competition, while introducing heterogeneity between groups leads to a significant intensification of conflict as well as increased volatility, thereby reducing earnings of contest participants and increasing inequality. We further find that heterogeneous groups share the labor much more equally than predicted by theory, and that in asymmetric contests group members change the way in which they condition their efforts on those of their peers. Implications for contest designers are discussed.

The Collective Action Problem but Not Numerical Superiority Explains Success in Intergroup Encounters in Verreaux’s Sifaka (Propithecus verreauxi): Implications for Individual Participation and Free-Riding

Socioecological theory predicts that larger groups are able to gain better access to resources because of their numerical advantage in intergroup contests. Nevertheless, the numerical superiority hypothesis has received mixed support, perhaps due to the collective action problem and sex differences in strategies and payoffs. We tested the numerical superiority and the collective action problem hypotheses using 141 intergroup encounters in a 7-year dataset on 5 groups of Verreaux’s sifaka (Propithecus verreauxi) at Ankoatsifaka Research Station in Kirindy Mitea National Park in western Madagascar. Contrary to the numerical superiority hypothesis, larger groups were not more likely to supplant smaller groups, nor was success determined by participation of a larger number of one sex. Consistent with a collective action problem, group size negatively affected participation. Notably, the sexes differed in which factors predicted individual participation. Male participation, which may serve to defend mating opportunities with resident females and signal competitive ability to neighboring groups, was greater than female participation, though we did not detect any strong predictor of male participation. Female participation was more likely during the lactation season, when food resources were scarce, than during the mating season. Contrary to previous studies, which argued that mothers with infants avoid intergroup conflicts because of the risk of infanticide, our study suggests that maternal energetic stress due to lactation and food scarcity increases the benefits of defending resources and instead drives mothers to participate in intergroup encounters. Thus, individual reproductive payoffs provide the private incentives needed to induce collective action.

Intragroup Behavioral Changes Following Intergroup Conflict in Mountain Gorillas (Gorilla beringei beringei)

Group-living animals face a number of threats from extragroup conspecifics: from individuals seeking mating opportunities to rival groups attempting to access limited resources. The consequences of intergroup interactions can therefore include loss of mates, increased energy expenditure, and injury or death. There is increasing evidence that aggressive intergroup interactions can affect subsequent intragroup behavior, and that such post-conflict behavior may be directly related to the threats posed by different opponent groups (e.g., familiar vs. unfamiliar). However, empirical evidence for changes in intragroup behavior following intergroup conflict in social animals is limited. We compared the proximity and behavior of group members before and after 84 aggressive intergroup interactions involving 14 study groups of wild, habituated mountain gorillas (Gorilla beringei beringei) in Volcanoes National Park, Rwanda. Intergroup conflict affected subsequent intragroup behavior in several ways. We found that immediate movement away from the opponent following intergroup conflict was less likely between familiar groups compared to unfamiliar groups. However, both winning and losing groups spent twice as much time moving post-conflict, and losing groups spent less time resting. There was an increase in female intragroup affiliative interactions and a decrease in male intragroup agonistic interactions post-conflict. These results demonstrate that such intergroup contests can influence intragroup dynamics beyond the immediate period of interaction.

Motivating political support with group-based rewards

To compete in political contests, politicians need their supporters to contribute resources. We investigate how politicians can best allocate group-based prizes on a contingent basis to incentivize individual members of a group to exert effort on their behalf. We contrast contingent contracts with tournament style contests (Lazear and Rosen (Rank-order tournaments as optimum labor contracts. J Political Econ 1981; 89(5): 841–864)) and characterize when tournaments that create intergroup contests for prizes elicit more contributions than contingent contracts. By tying the allocation of prizes to the observed level of effort made by each group, politicians ameliorate the collective action problem that is created by the fact that each individual’s effort has only a minimal impact on the overall political contest. We examine the relative performance and stability of intergroup tournaments as a function of the number of groups and asymmetric group size.

Sex Differences in Human Peer Relationships: A Primate’s-Eye View

Bonds formed by nonhuman animals can illuminate the structure of human relationships. In the juvenile period, primates of many species that are genetically similar to humans form sex-differentiated bonds in which females spend more time with female kin, and males spend more time with unrelated same-sex peers. Research with humans suggests a similar sex difference, with one notable addition: Beginning in middle childhood, male peer groups begin engaging in complex activities, including intergroup contests. This additional component of human peer relations resembles that of chimpanzees (Pan troglodytes), one of humans’ closest living genetic relatives. Cross-species and developmental evidence can aid in constructing a theory of human peer relations that differs by sex.

Not all Group Members are created Equal: Heterogeneous Abilities in Inter-group Contests

Competition between groups is ubiquitous in social and economic life, and groups are typically not created equal. Here we experimentally investigate the implications of this general observation on the unfolding of symmetric and asymmetric competition between groups that are either homogeneous or heterogeneous in the ability of their members to contribute to the success of the group. Our main finding is that, in contrast with a number of theoretical predictions, efforts in contests involving heterogeneous groups are higher than in contests involving only homogeneous groups, leading to reduced earnings (to contest participants) and increased inequality. This effect is particularly pronounced in asymmetric contests, where both homogeneous and heterogeneous groups increase their efforts. We find that asymmetry between groups changes the way group members condition their efforts on those of their peers. Implications for contest designers are discussed.

Home field advantage, not group size, predicts outcomes of intergroup conflicts in a social bird

Research on cooperatively breeding birds usually focuses on social dynamics within the breeding group, but conflict between groups can also affect individual fitness and the evolution of sociality. Here we investigate the causes and consequences of competition between groups of communally breeding greater anis, Crotophaga major , over a 10-year field study. Social groups were spatially clustered into loose aggregations that showed a moderate degree of reproductive synchrony. However, competition between neighbouring groups for nesting sites was intense, occasionally leading to wholesale destruction of a group's nesting attempt and abandonment of the site. We documented 18 cases in which a group's entire clutch of eggs was ejected from the nest during the laying or incubation period, often accompanied by behavioural observations of conflict with a neighbouring group. Clutch destruction typically occurred when two groups attempted to nest in close proximity on high-quality sites: nearest-neighbour distance and nest site type were the strongest predictors of clutch destruction. Surprisingly, group size did not predict whether or not a group's clutch would be destroyed, and small groups sometimes ousted larger groups. By contrast, ‘home field advantage’ did have a significant effect: groups that had previously nested on the site were more likely to destroy the clutches of newly established groups, and this effect increased with the number of years that the group had nested there. Together, these results support previous evidence that competition between groups for high-quality nesting sites is an important driver of communal breeding, and they highlight the importance of location and past history in determining the outcome of intergroup contests in social species.

Cross-Cultural Sex Differences in Post-Conflict Affiliation following Sports Matches

The nature of ancestral human social structure and the circumstances in which men or women tend to be more cooperative are subjects of intense debate. The male warrior hypothesis proposes that success in intergroup contests has been vital in human evolution and that men therefore must engage in maximally effective intragroup cooperation [1-3]. Post-conflict affiliation between opponents is further proposed to facilitate future cooperation [4], which has been demonstrated in non-human primates [5] and humans [6]. The sex that invests more in post-conflict affiliation, therefore, should cooperate more. Supportive evidence comes from chimpanzees, a close genetic relative to humans that also engages in male intergroup aggression [7]. Here we apply this principle to humans by testing the hypothesis that among members of a large community, following a conflict, males are predisposed to be more ready than females to repair their relationship via friendly contact. We took high-level sports matches as a proxy for intragroup conflict, because they occur within a large organization and constitutesemi-naturalistic, standardized, aggressive, and intense confrontations. Duration or frequency of peaceful physical contacts served as the measure of post-conflict affiliation because they are strongly associated with pro-social intentions [8, 9]. Across tennis, table tennis, badminton, and boxing, with participants from 44 countries, duration of post-conflict affiliation was longer for males than females. Our results indicate that unrelated human males are more predisposed than females to invest in a behavior, post-conflict affiliation, that is expected to facilitate future intragroup cooperation.

Effects of group size and contest location on the outcome and intensity of intergroup contests in wild blue monkeys

Intergroup contests determine access to resources and influence the evolution of group living in social species. Asymmetries in resource-holding potential and payoffs should influence the outcome and intensity of such contests. We evaluated predictors of contest outcome and intensity using data collected over 40 months from 6 groups of wild blue monkeys, Cercopithecus mitis. We found increased odds of winning when a group was larger and used the contest site more than its opponent, and when contests occurred closer to the group's home range centre while farther from the opponent's centre. However, a larger difference in group size (across five pairs of opposing groups) did not predict a greater proportion of contests won by the larger group. Some evidence suggested increased odds of a draw when group sizes were more similar. In addition, contests were longer and more aggressive when groups were more similar in size and when the contest site was similarly central in both groups' home ranges. Contests were also more aggressive when the opposing groups' use of the contest site was more similar. Overall, asymmetries in resource-holding potential (i.e. group size) and/or payoffs related to the contest's location influenced a group's competitive advantage, the likelihood of a draw and the intensity of intergroup contests. Although comparable data are limited, it seems clear that both types of asymmetries can play a role in determining the outcome and intensity of intergroup contests, and that the relative power of each may vary across species.

Individual participation in intergroup contests is mediated by numerical assessment strategies in black howler and tufted capuchin monkeys.

Asymmetries in resource-holding potential between opposing groups frequently determine outcomes of intergroup contests. Since both numerical superiority and high intergroup dominance rank may confer competitive advantages, group members should benefit from assessing the relative strength of rivals prior to engaging in defensive displays. However, differences in individual assessment may emerge when cost-benefit trade-offs differ among group members. We examine the influence of numerical superiority and intergroup dominance relationships on individual participation in intergroup encounters in black howler monkeys (Alouatta pigra) and tufted capuchin monkeys (Sapajus nigritus). Black howlers responded with longer vocal displays during encounters with neighbours with an equal number of resident males, while tufted capuchins increased their participation with increasing relative male group size. Within each species, males and females responded similarly to varying numerical odds, suggesting that despite pay-off asymmetries between males and females, both sexes were similarly influenced by numerical asymmetries in deciding to participate in collective group defence. Whereas the outcome of contests among tufted capuchins was determined by relative male group size, reflected in a pronounced intergroup dominance hierarchy, the absence of dominance relationships among black howler groups may have provoked prolonged vocal displays in order to assess rival groups with matching competitive abilities.

Patch Occupation Time Predicts Responses by Grey-Cheeked Mangabeys (Lophocebus albigena) to Real and Simulated Neighboring Groups

When two social groups are close enough to hear each other’s short-range vocalizations but not yet close enough to see each other, they can choose whether to approach to initiate a direct contest. In this study, I evaluated whether expected payoffs and group resource-holding potential affected the likelihood of approach toward a neighboring group during naturally occurring intergroup interactions and experimental playback trials by grey-cheeked mangabeys (Lophocebus albigena). Females approached calling groups at low rates, and only if males did so as well. Groups that had recently arrived (≤30 min) at the interaction location were more likely to approach than groups that were in the interaction location for >30 min. Site residency likely indicates the degree of short-term, local resource exploitation, and explains this pattern of patch defense. There was no evidence that mangabeys defended home range core or peripheral areas, that males defended female mates, or that infant defense affected the likelihood of approach. Surprisingly, resource-holding potential had no effect on patterns of contest initiation, but may influence other aspects of intergroup contests. The unexpected importance of payoff asymmetries, relative to asymmetries in resource-holding potential, points to the need for an expanded theoretical framework.

The impact of management incentives in intergroup contests

In intergroup contests a manager advises and motivates her group’s members. Her rewards often depend on the subsequent contest expenditure of the members. I test whether such incentives undermine the credibility and effectiveness of a manager’s efforts. In the different experimental treatments the managers either benefit from very high or low expenditure or get a predetermined payment. The results show that different management incentives shape the expenditure of the group members even if managers have an advisory role only. However, group members follow recommendations more closely if management compensation is not linked to contest expenditures.

Social Exclusion: More Important to Human Females Than Males

Theoretical models based on primate evidence suggest that social structure determines the costs and benefits of particular aggressive strategies. In humans, males more than females interact in groups of unrelated same-sex peers, and larger group size predicts success in inter-group contests. In marked contrast, human females form isolated one-on-one relationships with fewer instrumental benefits, so social exclusion constitutes a more useful strategy. If this model is accurate, then human social exclusion should be utilized by females more than males and females should be more sensitive to its occurrence. Here we present four studies supporting this model. In Study 1, using a computerized game with fictitious opponents, we demonstrate that females are more willing than males to socially exclude a temporary ally. In Study 2, females report more actual incidents of social exclusion than males do. In Study 3, females perceive cues revealing social exclusion more rapidly than males do. Finally, in Study 4, females’ heart rate increases more than males’ in response to social exclusion. Together, results indicate that social exclusion is a strategy well-tailored to human females’ social structure.

Fight tactics in wood ants: individuals in smaller groups fight harder but die faster

When social animals engage in inter-group contests, the outcome is determined by group sizes and individual masses, which together determine group resource-holding potential ('group RHP'). Individuals that perceive themselves as being in a group with high RHP may receive a motivational increase and increase their aggression levels. Alternatively, individuals in lower RHP groups may increase their aggression levels in an attempt to overcome the RHP deficit. We investigate how 'group RHP' influences agonistic tactics in red wood ants Formica rufa. Larger groups had higher total agonistic indices, but per capita agonistic indices were highest in the smallest groups, indicating that individuals in smaller groups fought harder. Agonistic indices were influenced by relative mean mass, focal group size, opponent group size and opponent group agonistic index. Focal group attrition rates decreased as focal group relative agonistic indices increased and there was a strong negative influence of relative mean mass. The highest focal attrition rates were received when opponent groups were numerically large and composed of large individuals. Thus, fight tactics in F. rufa seem to vary with both aspects of group RHP, group size and the individual attributes of group members, indicating that information on these are available to fighting ants.

Inter‐unit contests within a provisioned troop of Sichuan snub‐nosed monkeys (Rhinopithecus roxellana) in the Qinling Mountains, China

Numerical superiority does not always ensure victory in intergroup contests. Although group size is likely to determine the maximum resource holding potential (RHP) of a group, the realized RHP is the collective outcome of individual group members' choices about participation in any given contest. For any group member, the choice about participation should be based on the assessment of costs and benefits that are affected by both ecological and social factors. In this study, we studied inter-unit contests in a provisioned troop of Sichuan snub-nosed monkeys (Rhinopithecus roxellana). We spent 368 hr in contact with 9 one-male units sharing the same home range, during which we recorded 148 inter-unit contests at a provisioning site. Inter-unit contests always started as inter-individual contests. Contests escalated only when the two individuals were of different age-sex classes and one was an adult male. When a contest escalated, additional individuals were likely to get involved, and the outcome of the contest depended on unit members' choices about participation. The superiority in the number of participants rather than the superiority in unit size led to victory in inter-unit contests, given that the difference in unit size did not predict a difference in the number of participants. Unit members were more likely to support others in inter-unit contests in winter when food was sparse than in spring when food was abundant. In addition, unit members were more likely to support others in escalated contests than in those resulting in displacement, indicating that they tended to alter the outcome of a contest to gain immediate benefit. Although males initiated most inter-unit contests, a clear win-loss was most likely when females joined the fight. This sex difference may reflect the benefits to males vs. females of living in a multi-level society.