Pattern, Distribution, and Function of Greeting Behavior Among Black-and-White Colobus

Various species of primates engage in greeting, a ritualized pattern of nonaggressive behavior that usually occurs during a reunion. Black-and-white colobus (Colobus guereza) perform overhead mounting, mounting, and embracing behavior soon after an aggressive act and in nonagonistic situations. We studied the pattern, distribution, and function of the greeting behavior in 2 captive groups of black-and-white colobus. Overhead mounting was the most frequent pattern, accounting for >60% of all greetings (N=333). In nonagonistic situations, younger subordinate individuals greeted an older dominant individual more frequently than vice versa. A dominant male in a small multimale group frequently initiated contacts with adult females, though he was the oldest in the group. Conversely, the dominant male in a large 1-male group never greeted group members. Immediately after performing the greeting, the greeter groomed the recipient in more than half the cases. In the large group, greeting frequency correlates positively with the age difference between the pair; however, there is no correlation between the relatedness, affiliation frequency, or aggression frequency of a pair in either the small or large group. The results are consistent with the hypothesis that greeting behavior functions as a tension-reducing mechanism in nonagonistic situations. There is no evidence, however, that greeting functions to express social status or to attract a mate. In addition, the frequent greeting by the adult male in the small multimale group may indicate that individuals affirm social bonds via greeting behavior.     

Greeting,aggression, and coalitions between male baboons: Demographic correlates

Interactions of asymmetrical, symmetrical, and unreciprocated greeting, and episodes of aggression and coalitions observed between 18 males, members of a large multi-harem colony of baboons (Papio hamadryas, P. cynocephalus, and their hybrids) housed in the Madrid Zoo, were recorded throughout 11 time blocks between January 1983 and March 1985. The major question investigated was whether the demographic characteristics of the colony affected the frequency with which males used three behavioural strategies, namely, greeting, aggression, and coalitions. It was found that under demographic conditions eliciting more intense male-male competition, for example, when the ratio of oestrous cycles per adult male was lower, males exchanged frequent greetings, especially the symmetrical form (i.e. notifying) and solicited more coalitions, but did not show any significant change in rates of dyadic aggression. Male phenotype-related effects on rates of greeting and coalitions were found as well. Results are discussed in relation with the suggested role of greeting and of coalition behaviour as strategies used in the negotiation of changes in competitive social relationship (e.g. consort changeovers). These strategies are regarded as social tactics alternative to direct aggression.     

The functions of the "Greeting Ceremony" among male mantled howlers (Alouatta palliata) on Agaltepec Island, Mexico

Nonhuman primates use greeting behaviors as nonaggressive communicatory signals in multiple social contexts. Adult male mantled howlers (Alouatta palliata) perform a ritual greeting that has been associated with bond-strengthening functions. The aim of this study is to explore the greeting patterns of male howlers living on Agaltepec Island, Mexico. Specifically, we analyzed the relationships between greetings and several individual, relational, and contextual variables, such as the expression of affiliation and agonism, dominance rank, age, kinship relationships, spatial organization, activity patterns, and subgrouping patterns. Greetings were more frequent between males with closer dominance ranks. Among those dyads that greeted at least once, dominant males initiated greetings more frequently than less-dominant males. On the other hand, more greetings were observed when one of the participants had recently returned to a subgroup and during locomotion. On the basis of these results, we propose that on Agaltepec greetings are a conflict management mechanism used between males of similar ranks. The fission-fusion social system of this group of howlers allows males with conflicting interests to remain separated, and greetings may reduce tension during fusion events.     

Post‐Conflict Non‐Aggressive Behaviours may be neither Friendly nor Conciliatory: Conflict Management of Male Hamadryas Baboons

The relational model of conflict resolution predicts that after an aggressive conflict there should be a motivational shift from aggression to attraction. Most tests of the reconciliation hypothesis assume, however, that all non‐aggressive post‐conflict behaviours between former opponents are motivationally homogeneous and qualify as friendly reunions. In fact, although the hypothesis predicts an increased occurrence of friendly contacts after conflicts, in practice, however, post‐conflict reunions often include a mixture of contact and non‐contact behaviours. Most reconciliation studies either (often) assume a conciliatory function for post‐conflict reunions or (less often) test functional predictions. Finally, the valuable relationships hypothesis predicts that conciliatory rates should be relatively higher between friends and allies than between non‐friends/allies. In this paper, we use data on non‐aggressive interactions following conflicts between adult male hamadryas baboons that are neither friends nor allies to assess the implications of all these important but largely overlooked issues. The analyses of the rate and temporal relation of non‐contact greeting (NCTG) to anxiety‐related behaviours and side‐directed aggression as well as of the behaviours used during non‐aggressive interactions with male and female third‐parties suggest that the NCTG used by males after conflicts were neither motivationally friendly nor functionally conciliatory. We point out that the gestures exchanged during these post‐conflict NCTG can be interpreted as formalized signals of equal status and that the rate and form of the greetings used by male opponents are indicative of high relationship insecurity and incompatibility respectively. We conclude that although male hamadryas’ post‐conflict NCTG are not conciliatory they may serve to assess their opponents’ attitude and to negotiate the restoring of their pre‐conflict levels of peaceful but non‐amicable co‐existence.     

Social relationships and ritualized greetings in adult male baboons (Papio cynocephalus anubis)

Greetings involving exchanges of ritualized sexual gestures are a common form of interaction among adult male baboons, although relatively little attention has been paid to them. In this study, we investigate how greetings reflect important aspects of the male's social relationships, including dominance rank, age/residence status, and cooperative tendencies. The results are based on over 600 greetings among 12 adult males recorded during a 4-month study of a troop of wild olive baboons near Gilgil, Kenya. Four of the adult males were older, lower-ranking, long-term residents, which frequently formed coalitions to take estrous females away from the eight young, higher-ranking males. Virtually all dyads greeted: greetings occurred more than twice as often as other types of male-male interactions; and nearly all greetings occurred in a neutral context, in which there was no resource at stake. The percentage of greetings completed, the frequency with which different gestures were employed, and the roles adopted by each male varied significantly across old-old, old-young, and young-young dyads. Greetings between young adult males were often interrupted or actively resisted, consistent with their unstable and ambiguous dominance relationships. Greetings between old-old dyads were usually completed and appeared consistent with their cooperative relationships. One pair of old males formed a stable, reciprocal coalition against young males, and this pair's greetings showed remarkable symmetry of roles. Greetings, we hypothesize, function to allow males to negotiate important aspects of their relationships, including cooperation.     

Greeting movements among adult males in a colony of baboons:Papio hamadryas,P. cynocephalus and their hybrids

Seven greeting movements performed by four adult males belonging to a colony of baboons (Papio hamadryas, P. cynocephalus and their hybrids) were described. The hamadryas male has more number and more refined repertoire of movements than the yellow baboons, what would mean that the greeting movements possess a species-specific nature. This characteristic is expressed also by the existence of some inappropriate responses of the yellow baboons to the greeting of the hamadryas male. The differences in the greeting movements between both species would be explained as characteristic consequences of their social system organization. The hybrid male, having acquired a system organization similar to that of the hamadryas male, has acquired the same signal code, too, according to this system. The hybridization tendency would be adaptive in those colonies or troops with mixed species.     

Dominance relationships among male hamadryas baboons (Papio hamadryas)

The concept of dominance has been shown to be useful in describing and predicting social interactions in group-living animals. However, the dominance relationships among adult hamadryas males (Papio hamadryas) are poorly understood, and systematic data are missing from the literature. This study investigated dominance relationships among male hamadryas baboons by applying novel statistical techniques to systematic detailed data on agonistic interactions. We also analyzed the dominance relationship with male age and evaluated the association between dominance and access to mating partners (i.e. the number of adult females per one-male unit (OMU)), food resources (i.e. monopolization of feeding areas), and greeting interactions. The derived dominance indexes showed that, in general, leader males were dominant over followers, and that dominance ability did not correlate with male age. Individual dominance values were very close to each other, suggesting that dominance relationships among hamadryas males were not very rigid. In addition, dominance values were positively correlated with number of adult females per OMU but not with feeding priority. Finally, greeting interactions occurred more frequently between individuals with similar dominance values and with low levels of dominance decidedness. We suggest the need of further studies, especially in wild populations, to confirm our findings.     

The benefits of social capital: close social bonds among female baboons enhance offspring survival

Sociality has evolved in many animal taxa, but primates are unusual because they establish highly differentiated bonds with other group members. Such bonds are particularly pronounced among females in species like baboons, with female philopatry and male dispersal. These relationships seem to confer a number of short-term benefits on females, and sociality enhances infant survival in some populations. However, the long-term consequences of social bonds among adult females have not been well established. Here we provide the first direct evidence that social relationships among female baboons convey fitness benefits. In a group of free-ranging baboons, Papio cynocephalus ursinus, the offspring of females who formed strong social bonds with other females lived significantly longer than the offspring of females who formed weaker social bonds. These survival benefits were independent of maternal dominance rank and number of kin and extended into offspring adulthood. In particular, females who formed stronger bonds with their mothers and adult daughters experienced higher offspring survival rates than females who formed weaker bonds. For females lacking mothers or adult daughters, offspring survival was closely linked to bonds between maternal sisters. These results parallel those from human studies, which show that greater social integration is generally associated with reduced mortality and better physical and mental health, particularly for women.     

Function of Notification in Papio hamadryas

Notification is a form of ritualized greeting behavior performed by adult male hamadryas baboons (Papio hamadryas). Originally observed in the wild when harem leaders signaled the direction of travel to their followers on daily foraging marches, its true function is unknown. Notification involves the approach of an adult male baboon to another male, the presentation of his hindquarters, then retreat. Researchers have previously linked notification to aggression avoidance, appeasement, and the dominance-subordination relationship between males. We observed instances of notification over 6 mo among 8 adult males in a troop of 61 captive-bred hamadryas baboons at Paignton Zoo Environmental Park, Devon, UK. We compared observation sessions in which notification occurred with matched control periods. We classified different forms of notification according to the recipient’s response: presenting, mounting, touching of the buttocks or genitals, or ignoring. Results suggest that different forms of notification may have different functions, including submission, alliance, and peacekeeping but not general greeting, recruitment, or appeasement.     

Maintenance of pair bond during the non-reproductive season in a monogamous pipefish <Emphasis Type="Italic">Corythoichthys haematopterus</Emphasis>

During the non-reproductive season, male and female Corythoichthys haematopterus came to a particular site every early morning and performed a series of displays, called greetings, for a few minutes. After exchanging greetings, they had no further interactions during the daytime. This behavioural pattern was basically the same as that observed during the reproductive season. All greeting pairs bred in the following reproductive season if both members were present. These results indicate that mate fidelity of this pipefish is very high and that daily greetings lasting only a few minutes were enough to maintain pair bond all year around.     

Stronger social bonds do not always predict greater longevity in a gregarious primate

In group‐living species, individuals often have preferred affiliative social partners, with whom ties or bonds can confer advantages that correspond with greater fitness. For example, in adult female baboons and juvenile horses, individuals with stronger or more social ties experience greater survival. We used detailed behavioral and life history records to explore the relationship between tie quality and survival in a gregarious monkey (Cercopithecus mitis stuhlmanni), while controlling for dominance rank, group size, and life history strategy. We used Cox proportional hazards regressions to model the cumulative (multi‐year) and current (single‐year) relationships of social ties and the hazard of mortality in 83 wild adult females of known age, observed 2–8 years each (437 subject‐years) in eight social groups. The strength of bonds with close partners was associated with increased mortality risk under certain conditions: Females that had strong bonds with close partners that were inconsistent over multiple years had a higher risk of mortality than females adopting any other social strategy. Within a given year, females had a higher risk of death if they were strongly bonded with partners that changed from the previous year versus with partners that remained consistent. Dominance rank, number of adult female groupmates, and age at first reproduction did not predict the risk of death. This study demonstrates that costs and benefits of strong social bonds can be context‐dependent, relating to the consistency of social partners over time.     

Development in the Usage and Comprehension of Greeting Calls in a Free‐Ranging Group of Japanese Macaques (Macaca fuscata)

The main objective of this study is to clarify the developmental process involved in both the usage of greeting calls and the response to greeting calls by Japanese macaques. These greeting calls facilitate affiliative interactions by communicating benign intent. Specifically, individuals frequently emit greeting calls when interacting with less‐predictable individuals. Here, we examined whether the targets at which greeting calls are directed, along with associated behavioral responses, differed among the age classes, by conducting a cross‐sectional observation of females aged 0–5 yr and their mothers. We found that infant females showed a weak tendency to emit greeting calls at no specific receivers, unlike that by older females. Adult females emitted greeting calls more frequently when approaching unrelated females than related females. In contrast, young adult or juvenile females exhibited no significant difference in the proportion of the calls with related and unrelated conspecifics. Adult and young adult females were more likely to respond affiliatively to other individuals that approached using greeting calls compared with silent approaches, whereas juveniles did not exhibit different responses to the two types of approaches. This study showed that the target‐specific usage and affiliative response to greeting calls emerge with changes in the developmental stage. Furthermore, the fact that even young adults did not use greeting calls as adults indicates that the usage of greeting calls is modified in conjunction with the expansion of social relationships.     

Behavioral variation and reproductive success of male baboons (Papio anubis x Papio hamadryas) in a hybrid social group

We take advantage of an array of hybrid baboons (Papio anubis x Papio hamadryas) living in the same social group to explore the causes and consequences of different male mating strategies. Male hamadryas hold one-male units and exhibit a sustained, intense interest in adult females, regardless of the latter's reproductive state. Anubis baboons, by contrast, live in multi-male, multi-female groups where males compete for females only when the latter are estrous. These two taxa interbreed to form a hybrid zone in the Awash National Park, Ethiopia, where previous work has suggested that hybrid males have intermediate and ineffective behavior. Here, we first examine male mating strategies with respect to morphological and genetic measures of ancestry. We found significant relationships between behavioral measures and morphology; males with more hamadryas-like morphology had more hamadryas-like behavior. However, genetic ancestry was not related to behavior, and in both cases intermediates displayed a previously unreported level of behavioral variation. Furthermore, male behavior was unrelated to natal group. Second, we evaluated reproductive success by microsatellite-based paternity testing. The highest reproductive success was found for individuals exhibiting intermediate behaviors. Moreover, over nine years, some genetically and morphologically intermediate males had high reproductive success. We conclude that the behavior of hybrid males is therefore unlikely to be an absolute barrier to admixture in the region.     

Bonds of bros and brothers: Kinship and social bonding in postdispersal male macaques

Group‐living animals often maintain a few very close affiliative relationships—social bonds—that can buffer them against many of the inevitable costs of gregariousness. Kinship plays a central role in the development of such social bonds. The bulk of research on kin biases in sociality has focused on philopatric females, who typically live in deeply kin‐structured systems, with matrilineal dominance rank inheritance and life‐long familiarity between kin. Closely related males, in contrast, are usually not close in rank or familiar, which offers the opportunity to test the importance of kinship per se in the formation of social bonds. So far, however, kin biases in male social bonding have only been tested in philopatric males, where familiarity remains a confounding factor. Here, we studied bonds between male Assamese macaques, a species in which males disperse from their natal groups and in which male bonds are known to affect fitness. Combining extensive behavioural data on 43 adult males over a 10‐year period with DNA microsatellite relatedness analyses, we find that postdispersal males form stronger relationships with the few close kin available in the group than with the average nonkin. However, males form the majority of their bonds with nonkin and may choose nonkin over available close kin to bond with. Our results show that kinship facilitates bond formation, but is not a prerequisite for it, which suggests that strong bonds are not restricted to kin in male mammals and that animals cooperate for both direct and indirect fitness benefits.     

Food Competition and Linear Dominance Hierarchy Among Female Chimpanzees of the Taï National Park

Dominance rank in female chimpanzees correlates positively with reproductive success. Although a high rank obviously has an advantage for females, clear (linear) hierarchies in female chimpanzees have not been detected. Following the predictions of the socio-ecological model, the type of food competition should affect the dominance relationships among females. We investigated food competition and relationships among 11 adult female chimpanzees in the Taï National Park, Côte d'Ivoire (West Africa). We detected a formal linear dominance hierarchy among the females based on greeting behaviour directed from the subordinate to the dominant female. Females faced contest competition over food, and it increased when either the food was monopolizable or the number of competitors increased. Winning contests over food, but not age, was related to the dominance rank. Affiliative relationships among the females did not help to explain the absence of greetings in some dyads. However comparison post hoc among chimpanzee study sites made differences in the dominance relationships apparent. We discuss them based on social relationships among females, contest competition and predation. The cross-site comparison indicates that the differences in female dominance hierarchies among the chimpanzee study sites are affected by food competition, predation risk and observation time.     

Functional analysis of social staring behavior in an all-male group of mountain gorillas

A total 202 social staring episodes (prolonged gazing by one individual toward another within a short distance) were observed in various social contexts among six unrelated, adult, and subadult male mountain gorillas. Staring was not accompanied by distinct facial expressions by actors or recipients, irrespective of their age or dominance rank. Younger, subordinate animals tended to stare at elder, dominant animals more frequently than vice versa. Staring may have multiple functions depending on the social context. In the initiation of non-agonistic interactions, staring rarely occurred, but was very successful for younger males in eliciting play or homosexual interactions from older animals. Staring was also directed by younger males to older males for greeting or appeasement. It may possibly play a role in reducing the increased social tension that occurs during or after conflict and in averting the potential conflict among older males. Younger males occasionally supplanted older males by staring at feeding spots. This slow supplantation is similar to the phenomenon of food sharing achieved through begging behavior in chimpanzees and bonobos. A prolonged gaze including staring by subordinates towards dominants may characterize the frequent and prolonged face-to-face interactions in the African great apes, and contrasts with a frequent gaze aversion by subordinates towards dominants in macaques or baboons. A difference between gorillas and other apes is that, chimpanzees and bonobos make positive contact with each other through eye contact, while gorillas simply stare at another without physical contact in greeting or appeasement process. Staring may serve an effective strategy for younger male gorillas to intervene safely in olders' conflict and sometimes to suppress or inhibit olders' performance, but their non-agonistic contacts or supporting attacks may not contribute to the establishment or support of social bonds between them. It is possible that staring may be common among the African great apes and man, and that it has evolved as a tactics to mask the dominant/subordinate relationships between individuals with multiple functions.     

Significance of the social relationships of Efe Pygmy men in the Ituri forest,Zaire

Efe Pygmies in northeast Zaire exhibit features of social organization (small group size, flexible but predominantly patrilocal residence, close relations with kin, and pair bonds) that are common to many hunting and gathering societies. This study uses methods commonly employed in studies of nonhuman primates to investigate the associations of Efe men with other individuals as a function of their age, sex, and kinship, and it tests the hypothesis that Efe are patrilocal because hunting efficiency is improved by hunting with male relatives. The analyses of 376 hours of focal behavior observations on 16 Efe men and observations of 71 cooperative group hunts show that the majority of associations between Efe men were with other adult men. Men associated preferentially with kin over nonkin, and they associated with close kin more than with distant kin. Men's close relationships (what we call “Companionships” sensu Smuts' “Friendships” in baboons) were predominantly with other adult men, but each man who cohabited with a woman had his strongest Companionship by far with that woman. Neither the efficiency of hunts nor the hunting success of individual men were found to be related to the degree of relatedness of hunters nor the proportion of relatives on the hunt. Alternative hypotheses concerning the functional significance of patrilocality and male kin groups are considered. We conclude that strong affiliative bonds between male kin and between males and females may arise as much from the need for strong allies in the face of competitive social situations as from economic or ecological necessity.     

Relationships Between Adult Male and Maturing Mountain Gorillas (Gorilla beringei beringei) Persist Across Developmental Stages and Social Upheaval

Across the animal kingdom, long‐term social relationships outside the context of reproductive pair bonds are rare. However, they have been demonstrated in some mammals including primates, cetaceans, and social carnivores. The ontogeny of such relationships is likely to depend on the benefits individuals can gain by cultivating them. Previous studies demonstrated that young mountain gorillas (Gorilla beringei beringei) have strong relationships with adult males, but little is known about the longevity of these bonds. Here, we examine the temporal stability of proximity relationships between coresident adult male and maturing gorillas in the habituated population monitored by the Dian Fossey Gorilla Fund International's Karisoke Research Center in Volcanoes National Park, Rwanda. We used spatial proximity data to assess the strength of relationships between adult males and juveniles, and tracked these relationships as the juveniles matured into subadults (3–4 yr later; n = 229 dyads) and then young adults (7 yr later; n = 42 dyads). The proximity relationships of juveniles of both sexes predicted their proximity relationships with adult males in both subadulthood and young adulthood. However some young adult males who had lost their top adult male proximity partner from juvenilehood developed new relationships with older males that had risen in the dominance hierarchy. These data suggest that (1) kin selection may play a more important role in social relationships between potential fathers and adult female offspring than previously suspected, and (2) when maturing males' foremost adult male social partners remain available to them, their relationships can be stable past the age at which younger males become reproductive competitors.     

Social relationships of mountain baboons: Leadership and affiliation in a non-female-bonded monkey

Instead of close and differentiated relationship among adult females, the accepted norm for savanna baboons, groups of Drakensberg mountain baboons (Papio ursinus) showed strong affiliation of females towards a single male. The same male was usually the decision-making animal in controlling group movements. Lactating or pregnant females focused their grooming on this “leader” male, producing a radially patterned sociogram, as in the desert baboon (P. hamadryas); the leader male supported young animals in the group against aggression and protected them against external threats. Unlike typical savanna baboons, these mountain baboons rarely displayed approach-retreat or triadic interactions, and entirely lacked coalitions among adult females. Both groups studied were reproductively one-male; male-female relationships in one were like those in a unit of hamadryas male at his peak, while the other group resembled the unit of an old hamadryas male, who still led the group, with a male follower starting to build up a new unit and already monoplizing mating. In their mountain environment, where the low population density suggests conditions as harsh for baboons as in deserts, adults in these groups kept unusually large distances apart during ranging; kin tended to range apart, and spacing of adults was greatest at the end of the dry, winter season. These facts support the hypothesis that sparse food is responsible for convergence with hamadryas social organization. It is suggested that all baboons, though matrilocal, are better categorized as “cross-sex-bonded” than “female bonded”.     

A Papionin Multilevel Society as a Model for Hominin Social Evolution

Multilevel societies are unique in their ability to facilitate the maintenance of strong and consistent social bonds among some individuals while allowing separation among others, which may be especially important when social and sexual bonds carry significant and reliable benefits to individuals within social groups. Here we examine the importance of social and sexual bonds in the multilevel society of hamadryas baboons (Papio hamadryas) and apply these principles to social evolution in Plio-Pleistocene hominins. The behavior, adaptations, and socioecology of baboons (Papio spp.) have long been recognized as providing an important comparative sample to elucidate the processes of human evolution, and the social system of hamadryas baboons in particular shares even more similarities with humans than that of other baboons. Here we draw parallels between processes during the evolution of hamadryas social organization and those characterizing late Pliocene or early Pleistocene hominins, most likely Homo erectus. The higher costs of reproduction faced by female Homo erectus, exacerbated by an increased reliance on difficult to acquire, nutrient-dense foods, are commonly thought to have been alleviated by a strengthening of male–female bonds (via male provisioning and the evolution of monogamy) or by the assistance of older, postreproductive females (via grandmothering). We suggest that both of these social arrangements could have been present in Plio-Pleistocene hominins if we assume the development of a multilevel society such as that in hamadryas baboons. The evolution of a multilevel society thus underlies the adaptive potential for the complexity that we see in modern human social organization.