Social rejection,exclusion, and shunning among the Gombe chimpanzees

From the observations at Gombe over the past 23 years it appears that group punishment of deviant behavior through ostracism, as practiced in human groups, has not yet evolved in a truly sophisticated way in chimpanzee society. However, cases of “social rejection or exclusion” have been observed in three different behavioral contexts. Most frequently, a chimpanzee is the target of hostility as the result of competitive interaction within the community; in such cases, social cohesion counterbalances rejection, typically leading to integration within a relatively stable pattern of dominance and social interaction. The occasional departure of an individual who has been the target of aggression —like the withdrawal of Evered after repeated attacks by Figan and Faban—seems due to persistent hostility by a few males rather than general “ostracism” by the group as a whole. A second form of exclusion concerns outsiders found in the home range of the group: in these cases, hostility is more generalized, particularly in response to the attempt of an adult female with offspring to join the community. Finally, there are the rarely observed instances of shunning a group member whose behavior seems abnormal the social rejection of Pepe and Old Mr. McGregor after they suffered from polio.     

Ostracism as a social and biological phenomenon: An introduction

Ostracism refers to the general process of rejection and exclusion, observed in human groups and in many other species. It occurs as “shunning” in small homogenous groups like the Amish or as rejection among children. Ostracism in various forms is also deeply embedded in our own legal tradition, and is used in the formal and informal legal procedures of other cultures, used to maintain order, to punish deviance, and to increase social cohesion. Hence, it is plausible to hypothesize that human ostracism may have physiological substrates or biological functions in addition to cultural, moral, and legal dimensions. Biological research shows that human emotions (anger, fear, reassurance, self-confidence) involve responses of the limbic system as well as ideas or thoughts in the cerebral cortex and that human behavior continually integrates biological and cultural factors. The legal system expresses and channels human behavior. For this reason laws should be more effective if their functions complement (rather than ignore) the function of the behavior being regulated. To look at law as it affects human behavior in the light of the life sciences does not imply the intention of finding a universally valid “natural law,” akin to theological or ideological doctrine. On the contrary, we find that humans can form radically different social systems due to the plasticity of their behavior. However, an interdisciplinary analysis of ostracism as a common area of behavior, combining biology, law, and the social sciences can produce insights that none of these fields alone can provide. Such an approach should increase our understanding of human nature and the functions of law.     

Task-specific effects of ostracism on imitative fidelity in early childhood

This study examined task specific effects of third-party ostracism on imitative fidelity in early childhood (N = 96, 3–6-year-olds). Start- and end-states of action sequences were manipulated to examine the effects of priming third-party ostracism versus affiliation on children's imitation of instrumental (i.e., action sequence with a different start- and end-state) versus social convention (i.e., action sequence with an identical start- and end-state) tasks. Children's performance was coded for imitative fidelity and children's explanations for their behavior. As predicted, imitative fidelity was highest and social convention explanations were most common when primed with ostracism in the social convention task. The data are consistent with our proposal that imitation serves an affiliative function in response to the threat of ostracism, a response amplified for social conventions.     

Human Sexual Differences in the Use of Social Ostracism as a Competitive Tactic

An aspect of social systems that is similar between chimpanzees and humans is that males form larger groups than females do. Both chimpanzee and human studies suggest that large groups are costlier for females than for males, so females attempt to reduce group size. Social ostracism of female group members occurs in both species and may serve as a mechanism for group size reduction. We formed groups of female and male children to examine directly whether human females would be more likely than males to employ social ostracism. We asked 7 female and 7 male groups of 10-yr-old children to compose and perform a play about a topic of interest to them. Female groups engaged in social ostracism more than male groups did. Further, within female groups, cortisol levels remained higher for female perpetrators of social ostracism than for their victims, suggesting that social ostracism is costly. In contrast, more frequent 1:1 conflictual behavior occurred in male than in female groups, but cortisol was unrelated to frequencies of 1:1 conflicts. Our results support the theory that human females find groups aversive and seek to reduce their size via social ostracism. Coalitions minimize the risk of retaliation but may induce costs.     

Bioaltruism reconsidered

Altruistic behavior is often regarded as sociobiology's most central theoretical problem, but is it? Altruism in biology, bioaltruism, has many meanings, which can be grouped into two categories. The first I will callcommon bioaltruism. It is primarily of ethological relevance. The second,evolutionary bioaltruism, is a special category in evolutionary respects in that it may indeed pose a problem for evolutionary theory. These categories are logically independent. Moreover, both of them are logically different from altruism in its everyday psychological or moral sense. Sociobiological examples of bioaltruistic behavior concern bioaltruism in the first sense only, so the theoretical problem ‘altruism’ is supposed to pose, is indeed nothing but a theoretical problem and the bioaltruism that actually occurs has no evolutionary relevance. Nevertheless, evolutionary theory is relevant to our understanding of the possibility of common bioaltruism, and that possibility — even though bioaltruism is conceptually different from ethical altruism — is relevant for ethicists: it sheds light on what we can ask people to do or not to do.     

Breaking the Social–Non‐social Dichotomy: A Role for Reptiles in Vertebrate Social Behavior Research?

Although social behavior in vertebrates spans a continuum from solitary to highly social, taxa are often dichotomized as either ‘social’ or ‘non‐social’. We argue that this social dichotomy is overly simplistic, neglects the diversity of vertebrate social systems, impedes our understanding of the evolution of social behavior, and perpetuates the erroneous belief that one group—the reptiles—is primarily ‘non‐social’. This perspective essay highlights the diversity and complexity of reptile social systems, briefly reviews reasons for their historical neglect in research, and indicates how reptiles can contribute to our understanding of the evolution of vertebrate social behavior. Although a robust review of social behavior across vertebrates is lacking, the repeated evolution of social systems in multiple independent lineages enables investigation of the factors that promote shifts in vertebrate social behavior and the paraphyly of reptiles reinforces the need to understand reptile social behavior.     

Capital punishment in tribal Montenegro: Implications for law,biology, and theory of social control

Clan execution among feuding tribal societies is proposed as a significant precursor to modern law. This form of ostracism is examined with respect to its social control functions and the indigenous assumptions about behavior, genes and demography that guide the behavior. It is suggested that there are very close parallels between modern legal systems and nonliterate ones based on “self-help”, when capital punishment is used by normal decision groups in the name of “national security”.     

Neurological correlates of social behavior

Experimental evidence obtained over the past decade in nonhuman primates suggests that there are neural structures necessary for the maintenance of social bonds and affiliative behavior. These include the amygdaloid nuclei, which is the critical brain area, and two anatomically closely related cortical structures—the temporal pole and posterior medial orbital cortex. Bilateral ablation of any of the areas results in a syndrome that varies from a quantitative decrease in affiliative behavior in confined colonies to total social isolation in naturally free-ranging groups. Lesions of these areas in adult females are also incompatible with the maintenance of the maternal-infant bond, but operated infants thrive and are well cared for. Species-typical behavior will determine the response of group members to lesioned conspecifics, and may vary from attempts to reintegrate the affected subject to attack and ostracism.The amygdaloid nuclei are hypothesized to be essential to placing an emotional bias on sensory information; thus this brain area is sensitive to, and its function dependent on, the social/environmental context of ongoing behavior. Brain impairment per se, does not necessarily result in ostracism and may be compatible with maintenance of social bondings depending upon the neural structures involved, subject's affective state, communication ability, and species typical behaviors. Observations of brain-impaired humans closely parallel studies in nonhuman primates.     

Behavioral and physiological correlates of ostracism

A model of social interaction is outlined and used to interpret behavioral and physiological correlates of ostracism. Reported findings suggest that several physiological systems contribute to behaviors that are associated with ostracism and that these as well as other systems are involved in responses to being ostracized. The ideas discussed in this article suggest that future research should (1) identify persons who may be genetically predisposed to engage in behaviors associated with ostracism, and (2) identify persons who are physiologically vulnerable to being ostracized.     

Exit,Punishment and Rewards in Commons Dilemmas: An Experimental Study

Commons dilemmas are interaction situations where a common good is provided or exploited by a group of individuals so that optimal collective outcomes clash with private interests. Although in these situations, social norms and institutions exist that might help individuals to cooperate, little is known about the interaction effects between positive and negative incentives and exit options by individuals. We performed a modified public good game experiment to examine the effect of exit, rewards and punishment, as well as the interplay between exit and rewards and punishment. We found that punishment had a stronger effect than rewards on cooperation if considered by itself, whereas rewards had a stronger effect when combined with voluntary participation. This can be explained in terms of the ‘framing effect’, i.e., as the combination of exit and rewards might induce people to attach higher expected payoffs to cooperative strategies and expect better behaviour from others.     

A test of the haploid susceptibility hypothesis using a species with naturally occurring variation in ploidy

The haploid susceptibility hypothesis (HSH) was proposed as an explanation for how behavioral roles in haplodiploid social systems evolved. It posits that haploid males are more susceptible to disease than diploid females due to decreased genetic variability at key disease resistance loci. The resulting decreased immunocompetence is hypothesized to have played a role in the evolution of social behavior by limiting the behavioral repertoire haploids perform. Here, we test this hypothesis in a study system that separates ploidy from behavioral sex roles: Polistes dominulus, a social wasp, has colonies with naturally occurring diploid males. We report results from two immune function assays—hemolymph phenoloxidase activity and encapsulation response—performed on haploid males, diploid males, and diploid females. Our data suggest that ploidy is not a significant contributor to immune function in P. dominulus; thus, our data do not support the HSH for the evolution of behavioral roles. Instead, our data indicate that time of emergence is the best predictor of immune function in Polistes. We speculate that seasonal trends result from seasonal differences in pathogens and parasites.     

Evolution of social behavior by reciprocation

A general model is presented for the evolution of social behavior by reciprocation. The results of our model apply to social traits which are transmitted from one generation to the next by a process which guarantees that the frequency of the trait in one generation is directly related to its fitness in the preceding generation. The basic parameters of the model are α, the number of interactions per generation, and β, the number of these interactions which are with individuals who are perceived as strangers. It is shown that so long as α/β can be made large, social reciprocation may increase when arbitrarily rare even in the absence of population structure.This conclusion appears to be at odds with several recent investigations of Axelrod & Hamilton (1981) and Boorman & Levitt (1980). We use our model to reconcile these various approaches. By casting Axelrod & Hamilton's (1981) single-partner model in terms of the general parameters, α and β, we show that social reciprocation can increase when arbitrarily rare in a homogeneous population dominated by non-cooperators. Using a gene frequency approach, Boorman & Levitt (1973, 1980) demonstrated the existence of a selection threshold in frequency of the social trait, which must be surmounted for social reciprocation to increase. We show our analysis of reciprocation to be consistent with Boorman and Levitt's result, since for our general model the cost to the social individuals of learning the non-social's identity goes to zero as the ratio α/β gets large. We also use our general model to study two multi-partner models not considered elsewhere, which differ in regards to the memory capabilities assigned to the organism.Finally we use our model to compare directly the evolution of social behavior by reciprocation with the main alternate hypothesis, kin selection. We show that an act which accrues some cost ?c to the fitness of the donor while benefiting a recipient by b, will increase in frequency so long as c/b < Φ (equation (30)), where Φ is defined as the “coefficient of reciprocation” or probability that a cooperative act is reciprocated. By comparing the coefficient of reciprocation with the coefficient of relatedness of kin selection, direct comparisons of the two hypotheses may be made.     

The Effect of Ostracism and Optional Participation on the Evolution of Cooperation in the Voluntary Public Goods Game

Not only animals, plants and microbes but also humans cooperate in groups. The evolution of cooperation in a group is an evolutionary puzzle, because defectors always obtain a higher benefit than cooperators. When people participate in a group, they evaluate group member’s reputations and then decide whether to participate in it. In some groups, membership is open to all who are willing to participate in the group. In other groups, a candidate is excluded from membership if group members regard the candidate’s reputation as bad. We developed an evolutionary game model and investigated how participation in groups and ostracism influence the evolution of cooperation in groups when group members play the voluntary public goods game, by means of computer simulation. When group membership is open to all candidates and those candidates can decide whether to participate in a group, cooperation cannot be sustainable. However, cooperation is sustainable when a candidate cannot be a member unless all group members admit them to membership. Therefore, it is not participation in a group but rather ostracism, which functions as costless punishment on defectors, that is essential to sustain cooperation in the voluntary public goods game.     

Innovation in wild chimpanzees (Pan troglodytes)

Innovations of behavior have major implications for the concept of culture in animals. Innovation has rarefy been documented in wild animal populations. The chimpanzees of the Taï National Park, Côte d’Ivoire, spontaneously included new patterns of behavior in their repertoire during our study. Some innovations were incorporated into a minority of the group members’ repertoire, whereas others, such as building day nests in trees and on the ground more frequently, became general behaviors. Similarly, new contexts for use of leaf- clipping behavior and novel ways of feeding on some leaves appeared and rapidly became part of the behavior of most group members. The environmental parameters were too stable to explain these new forms of behavior, as most of them took place within 1 month during the same dry season. In a similar way, leaf- grooming acquired a new function in Gombe chimpanzees. A process similar to social conventions could explain the emergence of these new functions for an existing behavior in a way rather similar to human “fashion. ” Three major characteristics of human culture — the absence of individual variations in the performance of the behavior, the general use of the behavior by group members, and the ability to modify the function of a behavior — characterize leaf-clipping in Taï chimpanzees and leaf- grooming in Gombe chimpanzees.     

Maintaining cooperation in social-ecological systems:

Natural resources are vulnerable to over-exploitation in the absence of effective management. However, norms, enforced by social ostracism, can promote cooperation and increase stock biomass in common-pool resource systems. Unfortunately, the long-term sustainable use of a resource is not assured even if cooperation, maintained by ostracism and aimed at optimizing resource use, exists. Here, using the example of fisheries, we show that for a cooperative to be maintained by ostracism over time, it often must act inefficiently, choosing a ‘second-best’ strategy where the resource is over-harvested to some degree. Those cooperatives that aim for maximum sustainable profit, the “first-best” harvest strategy, are more vulnerable to invasion by independent harvesters, leading to larger declines in the fish population. In contrast, second-best strategies emphasize the resistance to invasion by independent harvesters over maximizing yield or profit. Ultimately, this leads to greater long-run payoffs to the resource users as well as higher resource stock levels. This highlights the value of pragmatism in the design of cooperative institutions for managing natural resources.     

Attachment Status Affects Heart Rate Responses to Experimental Ostracism in Inpatients with Depression

Depression is assumed to be both a risk factor for rejection and a result of it, and as such constitutes an important factor in rejection research. Attachment theory has been applied to understand psychological disorders, such as depression, and can explain individual differences in responses to rejection. Research on autonomic nervous system activity to rejection experiences has been contradictory, with opposing strings of argumentation (activating vs. numbing). We investigated autonomic nervous system-mediated peripheral physiological responses (heart rate) to experimentally manipulated ostracism (Cyberball) in 97 depressed patients with organized (n = 52) and disorganized attachment status (n = 45). Controlling for baseline mean heart rate levels, depressed patients with disorganized attachment status responded to ostracism with significantly higher increases in heart rate than depressed patients with organized attachment status (p = .029; ηp² = .051). These results suggest that attachment status may be a useful indicator of autonomic responses to perceived social threat, which in turn may affect the therapeutic process and the patient-therapist relationship.     

Hierarchical structure and the influence of individual attributes in the captive squirrel monkey (<Emphasis Type="Italic">Saimiri collinsi</Emphasis>)

The dominance structure of primate social groups varies widely. In addition to the groups’ composition, intrinsic attributes such as sex, body size and life experience are important factors that can affect hierarchical dominance relations. All primates are social animals, and the social environment has a direct influence on the physiological conditions of vital systems such as immunological, reproductive and cardiovascular systems. In this study, we analyze the hierarchical structure of Saimiri collinsi in captivity, including the hierarchical structure type, the influence of individual intrinsic characteristics (sex, age, weight and origin—born in captivity or in the wild) based on the prior-attributes model, the relation between agonistic behavior frequency and hierarchical position, and hierarchy steepness, which represents the dominance gradient. We found that the group order was characterized by a partial hierarchy: a dominance position could be occupied by more than one individual simultaneously, including individuals of both sexes. Intrinsic characteristics had no influence on hierarchical structure, with the exception of the male in the highest hierarchical position, which had a markedly larger body than all other group members. Thus, the prior-attributes model did not apply to hierarchical formation of S. collinsi in captivity. Only the frequency of agonistic behavior of males correlated with their hierarchical position, and they differed from all other group members in their more aggressive behavior. The steepness between adjacent positions along the dominance gradient was significant only between the dominant male and the next individual in the group, with a smooth gradient between the other positions in the rank. As the access to resources is directly related to hierarchical dominance, a smooth dominance gradient is to be expected in species that form very large groups, such as wild Saimiri populations.     

Fitness in stratified societies

When wealth or social status can be transmitted from parents to offspring and when fitness depends on wealth or social status, evolutionary consequences of individual transmission strategies can be described by a parameter, called long-term fitness by Rogers (1990), which is the expected relative contribution of an individual to the gene pool in the long-term future. We show how to measure and use this parameter in two models of general interest in sociobiology. First, we construct a system with social classes and hypergynous marriage. Our treatment includes a method for computing the fitnesses of the two sexes separately. As expected, upper-class males have the highest long-term fitness in this kind of social structure, followed by lower-class females, then lower-class males and upper-class females. Upper-class preference for sons would be favored by selection in this system, but not female unwillingness to marry down—in this sense such systems do not conform to a Darwinian model. We then study a system with one sex and three social classes, the poorest of which has very low single generation fitness. In this system, the class with the highest single generation fitness does not have the highest long-term fitness. We suggest that this system is a useful model for understanding the changes in reproductive behavior that occured during the demographic transition in Europe. We suggest that the absence of a destitute lower class in Africa may help explain the failure, so far, for signs of demographic transition to appear in Africa.     

Pukhtunwali: Ostracism and honor among the Pathan Hill tribes

The Pathan Hill tribes provide an example of the functional role of ostracism in a face- to-face, kin-based society. “Pukhtunwali,” the Code of the Pathans, regulates the uses of ostracism, as a response to the conflict between an individual's desire for freedom and the necessity of tribal unity. The most striking use of ostracism among the Pathans is the rejection by the tribe or clan of one of its own members whose behavior might lead to feud. If a member of a group has committed an act likely to provoke a reprisal, which may be directed against any individual of that group, the guilty person may be expelled. By ostracizing the person, the group is both punishing him, and withdrawing its support. In Pathan society ostracism functions simultaneously to deter behavior that violates customary legal norms, to punish specific acts that are culturally defined as improper, and to unify the primary reference group on which individuals depend for protection and economic support.     

Relatedness does not predict vigilance in a population of the social rodent Octodon degus

The possibility that social foragers adjust and coordinate their scanning activity when in the presence of close relatives to attain inclusive fitness benefits remains controversial and scarcely examined. To this aim, we first tested the null hypothesis of no association between foraging individuals of the diurnal rodent, Octodon degus and their pairwise relatedness (six microsatellite loci), under natural conditions. Secondly, we examined the influence of relatedness on scan effort (percent overlapping) and temporal distribution of scanning using linear regression. Finally, we evaluated whether temporal distributions of scanning were significantly lower (coordination) or higher (synchrony) than random expectations using bootstrapping. We found that pairwise relatedness between focal degus and their foraging partner did not influence the scan effort or the temporal distribution of scanning. These original, field-based findings imply that vigilance behavior in socially foraging degus is unlikely to be kin-selected and adds to results from previous lab studies in that kinship remains a poor predictor of social behavior in these animals. Overall, our study adds to others revealing that kin selection may not have had an impact on aspects of social behavior such as vigilance during social foraging.