Dominance style in female guerezas (Colobus guereza Rüppell 1835)

Socioecological models assume that quality and distribution of food ultimately determine female social relationships: a high quality diet and clumped food distribution should result in the establishment of a hierarchy with stable rank relations which is supported by empirical studies on frugivorous cercopithecines. By contrast, folivorous species with their low quality diet and dispersed food distribution should have egalitarian social relationships but empirical data are very rare. This study on female guerezas of a zoo group aimed to test the models in a colobine species and the results largely agreed with the predictions of the models: facial expressions, vocalizations, and gestures were not used for signalling dominance or subordination. Unritualized aggressions occurred frequently but were of low intensity, and interventions by third parties were never observed. Aggressions were exchanged bidirectionally and this was true also for food stealing and retreats. All this indicated the lack of established rank relations. Allogrooming was distributed rather equally and showed no kin bias. All these features characterize egalitarian social relationships and, hence, support the socioecological models.     

Coalitions and male–male behavior in <Emphasis Type="Italic">Alouatta palliata</Emphasis>

Coalitions influence the establishment and maintenance of social relationships among males in primate species. In this study, we compare the social behavior of males between two groups of Alouatta palliata: a group that was recently taken over by a coalition of two males (Mt), and a group that had a stable composition for at least 9 months (Rh). We predicted that coalition partners would be more cooperative and less competitive than dyads formed by immigrant and long-term resident males, and dyads formed by long-term resident males. Additionally, we predicted that these dyadic trends should be reflected in more competition and less cooperation in the group that was taken over. As predicted, the coalition partners of Mt showed the highest levels of cooperation among all dyads and the second lowest rate of agonism. Cooperation was higher in the group that had a stable composition. Results from this study suggest that the social relationships of male mantled howlers vary as a function of familiarity between males and that in the context of coalitionary takeovers, coalitionary males are highly cooperative. Cooperation is lower in groups recently taken over and competition is more intense, perhaps as a consequence of the process of establishment and reorganization of power relationships within some dyads. In the future, we must determine the frequency of coalitionary takeovers in this population and assess its ultimate consequences for male–male social relationships.     

Social learning in cooperative dilemmas

Helping is a cornerstone of social organization and commonplace in human societies. A major challenge for the evolutionary sciences is to explain how cooperation is maintained in large populations with high levels of migration, conditions under which cooperators can be exploited by selfish individuals. Cultural group selection models posit that such large-scale cooperation evolves via selection acting on populations among which behavioural variation is maintained by the cultural transmission of cooperative norms. These models assume that individuals acquire cooperative strategies via social learning. This assumption remains empirically untested. Here, I test this by investigating whether individuals employ conformist or payoff-biased learning in public goods games conducted in 14 villages of a forager–horticulturist society, the Pahari Korwa of India. Individuals did not show a clear tendency to conform or to be payoff-biased and are highly variable in their use of social learning. This variation is partly explained by both individual and village characteristics. The tendency to conform decreases and to be payoff-biased increases as the value of the modal contribution increases. These findings suggest that the use of social learning in cooperative dilemmas is contingent on individuals'' circumstances and environments, and question the existence of stably transmitted cultural norms of cooperation.     

The other cooperation problem: generating benefit

Understanding how cooperation evolves is central to explaining some core features of our biological world. Many important evolutionary events, such as the arrival of multicellularity or the origins of eusociality, are cooperative ventures between formerly solitary individuals. Explanations of the evolution of cooperation have primarily involved showing how cooperation can be maintained in the face of free-riding individuals whose success gradually undermines cooperation. In this paper I argue that there is a second, distinct, and less well explored, problem of cooperation that I call the generation of benefit. Focusing on how benefit is generated within a group poses a different problem: how is it that individuals in a group can (at least in principle) do better than those who remain solitary? I present several different ways that benefit may be generated, each with different implications for how cooperation might be initiated, how it might further evolve, and how it might interact with different ways of maintaining cooperation. I argue that in some cases of cooperation, the most important underlying “problem” of cooperation may be how to generate benefit, rather than how to reduce conflict or prevent free-riding.     

Evolution of Cooperation: Does Selfishness Restraint Lie within?

Traditional models of how cooperative strategies succeed in evolution have largely focused on social interactions among individuals and selection acting at kin and group levels. A recent study at the genetic level suggests that cooperation may also be promoted by the evolution of gene-trait relationships that limit the range of possible cheating mechanisms that can evolve.     

How chimpanzees solve collective action problems

We presented small groups of chimpanzees with two collective action situations, in which action was necessary for reward but there was a disincentive for individuals to act owing to the possibility of free-riding on the efforts of others. We found that in simpler scenarios (experiment 1) in which group size was small, there was a positive relationship between rank and action with more dominant individuals volunteering to act more often, particularly when the reward was less dispersed. Social tolerance also seemed to mediate action whereby higher tolerance levels within a group resulted in individuals of lower ranks sometimes acting and appropriating more of the reward. In more complex scenarios, when group size was larger and cooperation was necessary (experiment 2), overcoming the problem was more challenging. There was highly significant variability in the action rates of different individuals as well as between dyads, suggesting success was more greatly influenced by the individual personalities and personal relationships present in the group.     

The evolution of reciprocity in sizable groups

Recently, several authors have investigated the evolution of reciprocal altruism using the repeated prisoner's dilemma game. These models suggest that natural selection is likely to favor behavioral strategies leading to reciprocal cooperation when pairs of individuals interact repeatedly in potentially cooperative situations. Using the repeated n-person prisoner's dilemma game, we consider whether reciprocal altruism is also likely to evolve when social interactions involve more individuals. We show that the conditions that allow the evolution of reciprocal cooperation become extremely restrictive as group size increases.     

Fight for your breeding right: hierarchy re-establishment predicts aggression in a social queue

Social aggression is one of the most conspicuous features of animal societies, yet little is known about the causes of individual variation in aggression within social hierarchies. Recent theory suggests that when individuals form queues for breeding, variation in social aggression by non-breeding group members is related to their probability of inheriting breeding status. However, levels of aggression could also vary as a temporary response to changes in the hierarchy, with individuals becoming more aggressive as they ascend in rank, in order to re-establish dominance relationships. Using the group-living fish, Neolamprologus pulcher, we show that subordinates became more aggressive after they ascended in rank. Female ascenders exhibited more rapid increases in aggression than males, and the increased aggression was primarily directed towards group members of adjacent rather than non-adjacent rank, suggesting that social aggression was related to conflict over rank. Elevated aggression by ascenders was not sustained over time, there was no relationship between rank and aggression in stable groups, and aggression given by ascenders was not sex-biased. Together, these results suggest that the need to re-establish dominance relationships following rank ascension is an important determinant of variation in aggression in animal societies.     

Paternal Relatedness Predicts the Strength of Social Bonds among Female Rhesus Macaques

Forming strong, equitable, and enduring social bonds with a few individuals in a group carries adaptive benefits in terms of increased longevity, offspring survival and paternity success in birds and mammals, including humans. These recent insights generated a new interest in the factors creating variation in the strength of social relationships. Whether and how animals discriminate paternal kin from non-kin and bias their social behavior accordingly is being debated. This study explores the relative importance of dominance rank, age, maternal and paternal relatedness in shaping dyadic affiliative relationships in a group of 30 captive rhesus macaque females. The strength of social relationships, measured by the composite sociality index from observational data, was independently predicted in GLMMs by both maternal and paternal relatedness as well as rank similarity. In addition, social bonds between paternal half-sisters were stronger than between distantly related kin suggesting that females biased their affiliative effort towards paternal relatives. Despite identical relatedness coefficients bonds between mothers and their daughters were three times as strong as those between full sisters. Together these results add to the growing body of evidence for paternal kin biases in affiliative behavior and highlight that females prefer descendent over lateral kin.     

Avoiding the bullies: The resilience of cooperation among unequals

Can egalitarian norms or conventions survive the presence of dominant individuals who are ensured of victory in conflicts? We investigate the interaction of power asymmetry and partner choice in games of conflict over a contested resource. Previous models of cooperation do not include both power inequality and partner choice. Furthermore, models that do include power inequalities assume a static game where a bully’s advantage does not change. They have therefore not attempted to model complex and realistic properties of social interaction. Here, we introduce three models to study the emergence and resilience of cooperation among unequals when interaction is random, when individuals can choose their partners, and where power asymmetries dynamically depend on accumulated payoffs. We find that the ability to avoid bullies with higher competitive ability afforded by partner choice mostly restores cooperative conventions and that the competitive hierarchy never forms. Partner choice counteracts the hyper dominance of bullies who are isolated in the network and eliminates the need for others to coordinate in a coalition. When competitive ability dynamically depends on cumulative payoffs, complex cycles of coupled network-strategy-rank changes emerge. Effective collaborators gain popularity (and thus power), adopt aggressive behavior, get isolated, and ultimately lose power. Neither the network nor behavior converge to a stable equilibrium. Despite the instability of power dynamics, the cooperative convention in the population remains stable overall and long-term inequality is completely eliminated. The interaction between partner choice and dynamic power asymmetry is crucial for these results: without partner choice, bullies cannot be isolated, and without dynamic power asymmetry, bullies do not lose their power even when isolated. We analytically identify a single critical point that marks a phase transition in all three iterations of our models. This critical point is where the first individual breaks from the convention and cycles start to emerge.     

Social environment and sex differentiation in the false clown anemonefish, Amphiprion ocellaris

Plasticity in sex differentiation is known to be common in teleost fishes. Anemonefishes are protandrous; females are the largest and dominant members of social groups, displaying frequent aggressive behavior towards other members of groups. The second-ranked individuals become males and others remain as non-reproductive individuals. Here we examine the influence of social interaction in-group on sex differentiation in the false clown anemonefish, Amphiprion ocellaris, under laboratory conditions. Three juvenile anemonefish were kept in a tank for 180 days and their behaviors observed once a month. The social rank of individuals was distinguishable by their interactions in a group, with rank order clearly correlated with aggressive and appeasing behaviors. The dominant individuals occupied the shelter in the tank from the start to completion of the observation period. The body mass of dominant individuals increased compared to group-housed control fish, while third-ranked individuals showed growth suppression. The ratio of testicular tissue in gonads increased in dominant and second-ranked individuals but decreased in the third-ranked individuals. Differences in the plasma concentrations of estradiol, testosterone, and cortisol were not significant, but the concentration of 11-ketotestosterone was significantly higher in dominant individuals. These results suggest that, in false clown anemonefish, reproductive suppression of lower-ranked individuals becomes apparent in the first stage of group formation, and sex differentiation of upper-ranked individuals is gradually determined by long-term social interactions.     

Societal background influences social learning in cooperative decision making

Humans owe their ecological success to their great capacities for social learning and cooperation: learning from others helps individuals adjust to their environment and can promote cooperation in groups. Classic and recent studies indicate that the cultural organization of societies shapes the influence of social information on decision making and suggest that collectivist values (prioritizing the group relative to the individual) increase tendencies to conform to the majority. However, it is unknown whether and how societal background impacts social learning in cooperative interactions. Here we show that social learning in cooperative decision making systematically varies across two societies. We experimentally compare people's basic propensities for social learning in samples from a collectivist (China) and an individualist society (United Kingdom; total n?=?540) in a social dilemma and a coordination game. We demonstrate that Chinese participants base their cooperation decisions on information about their peers much more frequently than their British counterparts. Moreover, our results reveal remarkable societal differences in the type of peer information people consider. In contrast to the consensus view, Chinese participants tend to be substantially less majority-oriented than the British. While Chinese participants are inclined to adopt peer behavior that leads to higher payoffs, British participants tend to cooperate only if sufficiently many peers do so too. These results indicate that the basic processes underlying social transmission are not universal; rather, they vary with cultural conditions. As success-based learning is associated with selfish behavior and majority-based learning can help foster cooperation, our study suggests that in different societies social learning can play diverging roles in the emergence and maintenance of cooperation.     

Dynamics of alliance formation and the egalitarian revolution

Background

Arguably the most influential force in human history is the formation of social coalitions and alliances (i.e., long-lasting coalitions) and their impact on individual power. Understanding the dynamics of alliance formation and its consequences for biological, social, and cultural evolution is a formidable theoretical challenge. In most great ape species, coalitions occur at individual and group levels and among both kin and non-kin. Nonetheless, ape societies remain essentially hierarchical, and coalitions rarely weaken social inequality. In contrast, human hunter-gatherers show a remarkable tendency to egalitarianism, and human coalitions and alliances occur not only among individuals and groups, but also among groups of groups. These observations suggest that the evolutionary dynamics of human coalitions can only be understood in the context of social networks and cognitive evolution.

Methodology/Principal Findings

Here, we develop a stochastic model describing the emergence of networks of allies resulting from within-group competition for status or mates between individuals utilizing dyadic information. The model shows that alliances often emerge in a phase transition-like fashion if the group size, awareness, aggressiveness, and persuasiveness of individuals are large and the decay rate of individual affinities is small. With cultural inheritance of social networks, a single leveling alliance including all group members can emerge in several generations.

Conclusions/Significance

We propose a simple and flexible theoretical approach for studying the dynamics of alliance emergence applicable where game-theoretic methods are not practical. Our approach is both scalable and expandable. It is scalable in that it can be generalized to larger groups, or groups of groups. It is expandable in that it allows for inclusion of additional factors such as behavioral, genetic, social, and cultural features. Our results suggest that a rapid transition from a hierarchical society of great apes to an egalitarian society of hunter-gatherers (often referred to as “egalitarian revolution”) could indeed follow an increase in human cognitive abilities. The establishment of stable group-wide egalitarian alliances creates conditions promoting the origin of cultural norms favoring the group interests over those of individuals.     

Costly signaling and cooperation.

We propose an explanation of cooperation among unrelated members of a social group in which cooperation evolves because it constitutes an honest signal of the member's quality as a mate, coalition partner or competitor, and therefore results in advantageous alliances for those signaling in this manner. Our model is framed as a multi-player public goods game that involves no repeated or assortative interactions, so that non-cooperation would be a dominant strategy if there were no signaling benefits. We show that honest signaling of underlying quality by providing a public good to group members can be evolutionarily stable, and can proliferate in a population in which it is initially rare, provided that certain plausible conditions hold, including a link between group-beneficial signaling and underlying qualities of the signaler that would be of benefit to a potential mate or alliance partner. Our model applies to a range of cooperative interactions, including unconditionally sharing individually consumable resources, participating in group raiding or defense, and punishing free-riding or other violations of social norms.     

Evolution of equal division among unequal partners

One of the hallmarks of human fairness is its insensitivity to power: although strong individuals are often in a position to coerce weak individuals, fairness requires them to share the benefits of cooperation equally. The existence of such egalitarianism is poorly explained by current evolutionary models. We present a model based on cooperation and partner choice that can account for the emergence of a psychological disposition toward fairness, whatever the balance of power between the cooperative partners. We model the evolution of the division of a benefit in an interaction similar to an ultimatum game, in a population made up of individuals of variable strength. The model shows that strong individuals will not receive any advantage from their strength, instead having to share the benefits of cooperation equally with weak individuals at the evolutionary equilibrium, a result that is robust to variations in population size and the proportion of weak individuals. We discuss how this model suggests an explanation for why egalitarian behaviors toward everyone, including the weak, should be more likely to evolve in humans than in any other species.     

The economics of altruistic punishment and the maintenance of cooperation

Explaining the evolution and maintenance of cooperation among unrelated individuals is one of the fundamental problems in biology and the social sciences. Recent findings suggest that altruistic punishment is an important mechanism maintaining cooperation among humans. We experimentally explore the boundaries of altruistic punishment to maintain cooperation by varying both the cost and the impact of punishment, using an exceptionally extensive subject pool. Our results show that cooperation is only maintained if conditions for altruistic punishment are relatively favourable: low cost for the punisher and high impact on the punished. Our results indicate that punishment is strongly governed by its cost-to-impact ratio and that its effect on cooperation can be pinned down to one single variable: the threshold level of free-riding that goes unpunished. Additionally, actual pay-offs are the lowest when altruistic punishment maintains cooperation, because the pay-off destroyed through punishment exceeds the gains from increased cooperation. Our results are consistent with the interpretation that punishment decisions come from an amalgam of emotional response and cognitive cost-impact analysis and suggest that altruistic punishment alone can hardly maintain cooperation under multi-level natural selection. Uncovering the workings of altruistic punishment as has been done here is important because it helps predicting under which conditions altruistic punishment is expected to maintain cooperation.     

Negotiations over Grooming in Wild Vervet Monkeys (Chlorocebus pygerythrus)

Mutual grooming plays a central role in the establishment and maintenance of social relationships in primates. Allogrooming has two main functions: hygiene and bonding with partners. The duration of grooming bouts is commonly used in studies of the functional aspects of grooming, but few reflect on the proximate mechanisms that determine grooming bout lengths. As it is highly unlikely that groomer and groomee prefer exactly the same bout length, we are likely to observe the result of some form of negotiation. We currently lack information about the signals that primates employ to inform others about their intentions and desires concerning grooming interactions. From October 2006 until April 2007 we studied three behaviors shown in grooming interactions that could potentially have a signaling function in the negotiation process over the initiation and length of grooming bouts among adult females of two vervet groups freely ranging in the Loskop Dam Nature Reserve, South Africa: approaching another individual as far as that resulted in a grooming session, changing of the body position by the groomed individual, and lip smacking. We found that “approach” did not reliably predict which individual would receive grooming first, although approaching individuals groomed significantly more than those approached. Thus, in the context of grooming interactions, moving toward a group member may signal the willingness to invest. Body part presentations appeared to be the main signal used to demand a prolongation of the grooming by the partner. Finally, lip smacking was used under potentially stressful circumstances, notably shortly before using the mouth to groom the partner or an attempt to touch a mother’s infant. Our exploratory study hopefully inspires colleagues to start looking at the role of communication during cooperative interactions for a better appreciation of how animals manage cooperation and negotiate exchange rates.     

The Faith of Sacrifice: Leadership Trade-Offs in an Afro-Brazilian Religion

Despite secular trends in some countries, prestige-based authority in the form of religious leadership remains hugely influential in the everyday lives of millions of people around the world. Here, the costs and benefits of religious leadership are explored in an urban setting in northeastern Brazil. An economic game, within-group cooperation questionnaires, and social network analyses were carried out among adherents of an Afro-Brazilian religion. Results reveal that leaders display high levels of religious commitment and disproportionally provide cooperative services to group members. On the other hand, initiates cooperate less than leaders but do not differ in levels of received cooperation or social cohesion measures. This may indicate some level of exploitation or free-riding. Demographic and group variables also appear to play an important role in the degree of social cohesion a group achieves. These findings are discussed in the context of non-Western urban settings where religious leadership may represent both an alternative to social advancement and a crucial source of material aid, social support, and a strong sense of community.     

The evolutionary and ecological roots of human social organization

Social organization among human foragers is characterized by a three-generational system of resource provisioning within families, long-term pair-bonding between men and women, high levels of cooperation between kin and non-kin, and relatively egalitarian social relationships. In this paper, we suggest that these core features of human sociality result from the learning- and skill-intensive human foraging niche, which is distinguished by a late age-peak in caloric production, high complementarity between male and female inputs to offspring viability, high gains to cooperation in production and risk-reduction, and a lack of economically defensible resources. We present an explanatory framework for understanding variation in social organization across human societies, highlighting the interactive effects of four key ecological and economic variables: (i) the role of skill in resource production; (ii) the degree of complementarity in male and female inputs into production; (iii) economies of scale in cooperative production and competition; and (iv) the economic defensibility of physical inputs into production. Finally, we apply this framework to understanding variation in social and political organization across foraging, horticulturalist, pastoralist and agriculturalist societies.