Female Genito-genital Rubbing in a Group of Captive Chimpanzees

Wild and captive female bonobos exhibit a form of noncopulatory sexual behavior—genito-genital (GG) rubbing—in which 2 individuals rub their genital regions together. GG-rubbing in bonobos occurs in several contexts and may serve several functions, including tension reduction and reconciliation. It has not been reported for chimpanzees. In a study of captive, adolescent chimpanzees at the New Iberia Research Center, I observed that females rubbed genital regions at rates equivalent to those of bonobos, though the form of the behavior differed in significant ways from classic bonobo GG-rubbing. I describe the pattern of GG-rubbing in the chimpanzees and provide preliminary tests of 5 hypotheses for why the behavior occurs. All 4 females in the study group participated in GG-rubbing, though the frequency with which they initiated and participated varied. Females that GG-rubbed the most also groomed each other the most, supporting the hypothesis that GG-rubbing reinforces or at least reflects social bonds. The data do not support the hypothesis that females GG-rub to reconcile conflicts, to reduce tension during feeding, to signal social status, or to attract mates.     

Methodological and Evolutionary Aspects of Reconciliation among Primates

Affinitive contact between former opponents soon after a conflict has been demonstrated in a growing number of primate species. Several recent studies show that such contact reduces the probability of future conflicts, allows the recipient of aggression to reduce its anxiety, and restores tolerance between former opponents. Hence, these contacts can be termed reconciliation. In this paper, we critically discuss common methodological problems of studying reconciliation, examine functional aspects, and evaluate the existing variation in primate reconciliation in light of predictions derived from four untested hypotheses about its evolutionary origins. We find that the occurrence of reconciliation in primates is not limited to anthropoids. Neither is it limited to species with formalized dominance relations. Reconciliation is also not a prerequisite for life in permanent social groups. Instead, several lines of evidence support the hypothesis that reconciliation serves to maintain valuable social relationships between individuals. We suggest several more specific versions of this hypothesis and discuss reconciliation between kin, mates, and alliance partners, as well as a number of open questions pertaining to the mechanisms, functions and origins of reconciliation among primates.     

Bonobos Respond to Distress in Others: Consolation across the Age Spectrum

How animals respond to conflict provides key insights into the evolution of socio-cognitive and emotional capacities. Evidence from apes has shown that, after social conflicts, bystanders approach victims of aggression to offer stress-alleviating contact behavior, a phenomenon known as consolation. This other-orientated behavior depends on sensitivity to the other''s emotional state, whereby the consoler acts to ameliorate the other''s situation. We examined post-conflict interactions in bonobos (Pan paniscus) to identify the determinants of consolation and reconciliation. Thirty-six semi-free bonobos of all ages were observed at the Lola ya Bonobo Sanctuary, DR Congo, using standardized Post-conflict/Matched Control methods. Across age and sex classes, bonobos consoled victims and reconciled after conflicts using a suite of affiliative and socio-sexual behaviors including embracing, touching, and mounting. Juveniles were more likely to console than adults, challenging the assumption that comfort-giving rests on advanced cognitive mechanisms that emerge only with age. Mother-reared individuals were more likely to console than orphans, highlighting the role of rearing in emotional development. Consistent with previous studies, bystanders were more likely to console relatives or closely bonded partners. Effects of kinship, affiliation and rearing were similarly indicated in patterns of reconciliation. Nearby bystanders were significantly more likely to contact victims than more distal ones, and consolation was more likely in non-food contexts than during feeding. The results did not provide convincing evidence that bystander contacts served for self-protection or as substitutes for reconciliation. Overall, results indicate that a suite of social, developmental and contextual factors underlie consolation and reconciliation in bonobos and that a sensitivity to the emotions of others and the ability to provide appropriate consolatory behaviors emerges early in development.     

Nonconceptive Sexual Behavior in Bonobos and Capuchins

Sexual behavior by infecundable females, and by same-sex and adult-immature dyads, occurs in wild and captive bonobos (Pan paniscus). Proposed functions of these behaviors, in social primates generally, include practice, paternity confusion, exchange, and communication as well as appeasement. We used this framework to interpret and to compare observations of sexual behavior in a captive bonobo group and a wild white-faced capuchin (Cebus capucinus) group. In both species, (a) sexual behavior was no more frequent in cycling females than in pregnant or lactating females and (b) same-sex and adult-immature dyads engaged in as much mounting or genitogenital contact as adult heterosexual dyads did. The species differed in that (a) bonobos engaged in sexual behavior 65 times as frequently as capuchins, (b) only bonobos engaged in sexual contact other than ventrodorsal mounting during focal observation, and (c) bonobo sexual contact was concentrated most heavily in socially tense situations in adult female–female dyads, whereas capuchin sexual contact was concentrated most heavily in socially tense situations in adult male–male dyads. These data and published literature indicate that (a) practice sex occurs in both species, (b) paternity confusion may be a current function of C. capucinus nonconceptive sex, (c) exchange sex remains undemonstrated in capuchins, and (d) communication sex is more important to members of the transferring sex—female bonobos and male capuchins—than to members of the philopatric sex.     

Reconciling conflicts in a one-male society: the case of geladas (<Emphasis Type="Italic">Theropithecus gelada</Emphasis>)

Animals derive benefits from living in social groups but sociality also has its costs in that animals must compete with others for resources and mating opportunities. To cope with the conflict aftermath and social damage caused by competitive aggression, several group-living species use a variety of peace-keeping strategies. The affinitive post-conflict reunion of former opponents, defined as reconciliation, is the primary peace-keeping mechanism. In this study, we provide evidence for the occurrence of reconciliation and test some hypotheses on this post-conflict mechanism in geladas (Theropithecus gelada), a species often neglected in the study of post-conflict dynamics. The conciliatory contacts were uniformly distributed across the different sex–class combinations. Different from baboons, geladas did not show any particular kind of affinitive reconciliation behaviour. Notwithstanding the presence of a linear hierarchy, the dominance relationships did not affect the reconciliation dynamics. According to the valuable relationship hypothesis, coalitionary support seems to be a good predictor for a high level of conciliatory contacts. Finally, at an immediate level reconciliation plays a role in reducing renewed attacks by aggressors, which sought conciliatory contact more frequently than victims. In conclusion, even though the study of post-conflict behaviour in geladas needs to be continued, the patchy nature of their social network is a good model for testing some of the theoretical assumptions about primate conflict resolution.     

Post-conflict Social Events in Wild Mountain Gorillas (Mammalia,Hominoidea). I. Social Interactions between Opponents

Researchers have documented elevated rates of affinitive interaction between opponents shortly after aggressive conflicts, or reconciliation, in many primate species. Reconciliation may ameliorate the immediate negative effects of aggression by reducing the chance of further aggression between opponents and thus reducing tension, and may avert or repair damage to long-term social relationships important to the animals' fitness. Data on post-conflict interactions between opponents in two groups of wild mountain gorillas (Gorilla gorilla beringei) fail to show reconciliation after conflicts between females, between males, or between immatures, but females seem to commonly reconcile themselves with males after receipt of male aggression. Females and subordinate males often avoid same-sex opponents after conflicts. Females commonly retaliate against female aggressors, and post-conflict rates of aggression between females are higher than baseline levels. Females may not need to achieve reconciliation with each other because relationships between co-resident relatives are resiliant, while those between non-relatives are mostly neutral to antagonistic. Males are, however, important social partners and protectors of females, and female transfer is common. Thus, the results strongly support the ‘important relationships’ hypothesis for the function of reconciliation.     

Welcome Back: Responses of Female Bonobos (Pan paniscus) to Fusions

In species with a high degree of fission-fusion social dynamics, fusions may trigger social conflict and thus provide an opportunity to identify sources of social tension and mechanisms related to its alleviation. We characterized behavioral and endocrine responses of captive female bonobos (Pan paniscus) to fusions within a zoo facility designed to simulate naturalistic fission-fusion social dynamics. We compared urinary cortisol levels and frequencies of aggression, grooming and socio-sexual interactions between female bonobos while in stable sub-groups and when one “joiner” was reunited with the “residents” of another sub-group. We hypothesized that fusions would trigger increases in aggression and cortisol levels among reunited joiners and resident females. We further predicted that females who face more uncertainty in their social interactions following fusions may use grooming and/or socio-sexual behavior to reduce social tension and aggression. The only aggression on reunion days occurred between reunited females, but frequencies of aggression remained low across non-reunion and reunion days, and there was no effect of fusions on cortisol levels. Fusions did not influence patterns of grooming, but there were increases in socio-sexual solicitations and socio-sexual interactions between joiners and resident females. Joiners who had been separated from residents for longer received the most solicitations, but were also more selective in their acceptance of solicitations and preferred to have socio-sexual interactions with higher-ranking residents. Our results suggest that socio-sexual interactions play a role in reintegrating female bonobos into social groups following fusions. In addition, females who receive a high number of solicitations are able to gain more control over their socio-sexual interactions and may use socio-sexual interactions for other purposes, such as to enhance their social standing.     

Female bonobos use copulation calls as social signals

During mating events, females of many primate species produce loud and distinct vocalizations known as 'copulation calls'. The adaptive significance of these signals is considered to be in promoting the caller's direct reproductive success. Here, we investigated copulation calling in bonobos (Pan paniscus), a species in which females produce these vocalizations during sexual interactions with partners of both sexes. Females were more likely to call when mating with males than with females. We also observed a positive relationship between the likelihood of calling and partner rank, regardless of partner sex. Sexual activity generally increased with swelling size (an indicator of reproductive state) and, during their peak swelling, females called more with male than with female partners. Female bonobos are unusual among the non-human primates in terms of their heightened socio-sexuality. Our results suggest that in this species, copulation calls have undergone an evolutionary transition from a purely reproductive to a more general social function, reflecting the intrinsic evolutionary links between vocal behaviour and social cognition.     

Reconciliation and consolation in captive bonobos (Pan paniscus)

Although reconciliation in bonobos (Pan paniscus) has previously been described, it has not been analyzed heretofore by the postconflict (PC) match-control (MC) method. Furthermore, although reconciliation has been investigated before in this species, consolation has not. In this study we analyzed agonistic and affiliative contacts in all sex-class combinations to clarify and reevaluate the occurrence of reconciliation in bonobos via the PC-MC method. We also investigated the occurrence of consolation by analyzing the victims' triadic contact tendency (TCT), the influence of the sex of victims, and the relative occurrence of consolation and reconciliation. We collected 167 pairs of PC-MC observations in a captive group of bonobos (in Apeldoorn, The Netherlands). The conciliatory tendency (CCT) we obtained was tendentially lower than the mean value previously found for Yerkes captive chimpanzees. Close relationships, which were present in all female-female (FF) and some male-female (MF) dyads, positively affected reconciliation rates. When only adult PC-MC pairs (157) were considered, the mean TCTs and CCTs did not differ significantly. When we focused on types of PC affiliative contact, in the case of consolation we found a striking preference for sociosexual patterns. As to the relative occurrence of consolation and reconciliation, the highest level of the former was found in the absence of the latter. When reconciliation took place, consolation generally preceded it, suggesting that consolation may be a substitutive behavior. Our findings suggest that even if reconciliation remains the best option, consolation may be an alternative substitute for reconciliation that is used to buffer the tension originating from an unresolved conflict. Reconciliation and consolation are complex phenomena that are probably related to the life history of a group. Given that few studies have been conducted on this subject, we can not at this time make any generalizations regarding conflict resolution in certain species by comparing results among studies.     

Reconciliation and relationship quality in Assamese macaques (Macaca assamensis)

A consistent conclusion in reconciliation research is that animals that reconcile are likely to have strong social bonds. This has led to the hypothesis that reconciliation occurs most often between valuable social partners. We tested this hypothesis in a group of Assamese macaques (Macaca assamensis) living near a temple in Assam, India. Using focal sample and ad libitum data collection, we recorded the occurrence of reconciliation, grooming, and agonistic aiding, and the outcomes of approach. We used matrix association methods (TauKr correlation) to correlate reconciliation with grooming, aiding, and approach outcome. Females reconciled more often with females with which they had stronger grooming and aiding relationships. The correlation between reconciliation and aiding was significant for support to the aggressor and the victim. In contrast, no such correlations with reconciliation were found for males. This study provides evidence that females reconcile most often with valuable and compatible social partners. The results do not support the relationship-quality hypothesis for males, and we suggest that future studies give more consideration to the possibility that males reconcile for reasons other than to repair relationships with valuable partners.     

Tolerance allows bonobos to outperform chimpanzees on a cooperative task

To understand constraints on the evolution of cooperation, we compared the ability of bonobos and chimpanzees to cooperatively solve a food-retrieval problem. We addressed two hypotheses. The "emotional-reactivity hypothesis" predicts that bonobos will cooperate more successfully because tolerance levels are higher in bonobos. This prediction is inspired by studies of domesticated animals; such studies suggest that selection on emotional reactivity can influence the ability to solve social problems [1, 2]. In contrast, the "hunting hypothesis" predicts that chimpanzees will cooperate more successfully because only chimpanzees have been reported to cooperatively hunt in the wild [3-5]. We indexed emotional reactivity by measuring social tolerance while the animals were cofeeding and found that bonobos were more tolerant of cofeeding than chimpanzees. In addition, during cofeeding tests only bonobos exhibited socio-sexual behavior, and they played more. When presented with a task of retrieving food that was difficult to monopolize, bonobos and chimpanzees were equally cooperative. However, when the food reward was highly monopolizable, bonobos were more successful than chimpanzees at cooperating to retrieve it. These results support the emotional-reactivity hypothesis. Selection on temperament may in part explain the variance in cooperative ability across species, including hominoids.     

What Females Tell Males About Their Reproductive Status: Are Morphological and Behavioural Cues Reliable Signals of Ovulation in Bonobos (Pan paniscus)?

In many Old World primate species, female attractivity increases during the tumescent phase of the sexual swelling for a period that lasts considerably longer than oestrus‐related attractivity in other mammals. We examined the reliability of the swelling as an indicator of ovulation in captive bonobos, a species with a long and variable phase of maximum tumescence. Using a combined approach of (1) observations of sexual behaviour, (2) visual scoring of the sexual swelling and (3) analysis of faecal progestin to assess the timing of ovulation during 23 ovulatory cycles of eight adult females, we found that in 30% of these cycles the presumed day of ovulation did not fall within the period of maximum tumescence. When ovulation did occur during maximum swelling, it was more closely related to the end rather than the onset of the maximum swelling period. However, the pattern of sexual swelling was not a reliable indicator of ovulation. In addition, sexual behaviour of both sexes increased in frequency with the degree of the swelling but not around the time of ovulation. We conclude that swellings in bonobos provide honest information on the probability of ovulation, but not its exact timing, and that therefore the `obvious ovulation'‐hypothesis cannot explain the function of sexual swellings in bonobos.     

Factors underlying party size differences between chimpanzees and bonobos: a review and hypotheses for future study

Differences in party size and cohesiveness among females have been primary topics in socio-ecological comparisons of chimpanzees (Pan troglodytes) and bonobos (Pan paniscus). This paper aims to review previous studies that attempted to explain these differences and propose some hypotheses to be tested in future studies. Comparisons of recent data show that relative party size (expressed as a percentage of total group size) is significantly larger for bonobos than chimpanzees. Although the prolonged estrus of females, close association between mother and adult sons, female social relationships including unique homosexual behavior, and high female social status might be related to the increased party size and female cohesiveness of bonobos, these social and behavioral factors alone do not appear to explain the differences between the two species. Differences in ecological factors, including fruit-patch size, density of terrestrial herbs, and the availability of scattered foods that animals forage as they travel between large fruit patches could also contribute to the differences between chimpanzees and bonobos. However, these factors cannot fully account for the increased party size and female cohesiveness of bonobos. The higher female cohesiveness in bonobos may be explained by socio-ecological systems that reduce the cost in feeding efficiency incurred by attending mixed-sex parties. These systems may include female initiatives for party ranging movements as well as the factors mentioned above. Because of their geographical isolation, the two species probably evolved different social systems. Chimpanzees, whose habitats became very dry during some periods in the Pleistocene, likely evolved more flexible fission–fusion social systems to cope with seasonal and annual variation in food availability. On the other hand, bonobos had a large refugia forest in the middle of their range even during the driest periods in the Pleistocene. Therefore bonobos, whose habitats had more abundant food and smaller variation in food availability, probably evolved systems that help females stay in mixed parties without incurring large costs from contest and scramble competition.     

Intrasexual mounting in the beetle Diaprepes abbreviatus (L.).

The weevil Diaprepes abbreviatus shows three kinds of same-sex mountings: males mount other unpaired males, males mount males already engaged in copulation and females mount other females. Four hypotheses were evaluated in order to explain same-sex matings by males: (i) female mimicry by inferior males, (ii) dominance of larger males which affects the behaviour of small males, (iii) sperm transfer in which smaller males gain some reproductive success by 'hitchhiking' their sperm with the sperm of larger males, and (iv) poor sex recognition. Data from mate choice and sperm competition experiments rejected the female mimicry, dominance and sperm transfer hypotheses and supported the poor sex recognition hypothesis. We tested three hypotheses in order to explain female mounting behaviour: (i) females mimic male behaviour in order to reduce sexual harassment by males, (ii) females mount other females in order to appear larger and thereby attract more and larger males for mating, and (iii) female mimicry of males. The results of our mate choice experiments suggested that the female mimicry of males hypothesis best explains the observed female mounting behaviour. This result is also consistent with the poor sex recognition hypothesis which is the most likely explanation for male and female intrasexual mating behaviour in many insect species.     

The relationship between socio‐sexual behavior and salivary cortisol in bonobos: tests of the tension regulation hypothesis

Bonobos are known for their pacifistic behavior and their large repertoire of behaviors that are thought to serve conflict resolution. One is an unusual form of ventro‐ventral mounting that facilitates genital contacts (GC). Various hypotheses have been proposed to explain its function. In this study we tested predictions of the tension regulation hypothesis using salivary cortisol as a marker for social stress. The results indicate a temporal relationship between GC and cortisol levels. Compared with baseline data and matched samples of unrestricted food access, rates of GC increased when access to food sources was restricted. Cortisol levels were highest when access to food was constrained. However, because the behavioral and hormonal responses occurred when viewing the stimulus at a distance and preceded the physical presence of the stimulus, we conclude that the anticipation of a competitive situation was sufficient to induce social stress. Contrary to our prediction, targets of aggression did not have higher rates of GC nor did they solicit GC more often than others. Furthermore, higher GC rates did not correlate with a more pronounced decrease in cortisol levels. Not all results obtained in this study supported the predictions concerning the regulatory function of GC on social tension and further research is needed to explore this question. However, the results indicate that the anticipation of competition may be sufficient to induce a costly physiological response, and that high levels of resource competition may have lasting effects on physical stress and stress management. Am. J. Primatol. 71:223–232, 2009. © 2008 Wiley‐Liss, Inc.     

In Bonobos Yawn Contagion Is Higher among Kin and Friends

In humans, the distribution of yawn contagion is shaped by social closeness with strongly bonded pairs showing higher levels of contagion than weakly bonded pairs. This ethological finding led the authors to hypothesize that the phenomenon of yawn contagion may be the result of certain empathic abilities, although in their most basal form. Here, for the first time, we show the capacity of bonobos (Pan paniscus) to respond to yawns of conspecifics. Bonobos spontaneously yawned more frequently during resting/relaxing compared to social tension periods. The results show that yawn contagion was context independent suggesting that the probability of yawning after observing others'' yawns is not affected by the propensity to engage in spontaneous yawns. As it occurs in humans, in bonobos the yawing response mostly occurred within the first minute after the perception of the stimulus. Finally, via a Linear Mixed Model we tested the effect of different variables (e.g., sex, rank, relationship quality) on yawn contagion, which increased when subjects were strongly bonded and when the triggering subject was a female. The importance of social bonding in shaping yawn contagion in bonobos, as it occurs in humans, is consistent with the hypothesis that empathy may play a role in the modulation of this phenomenon in both species. The higher frequency of yawn contagion in presence of a female as a triggering subject supports the hypothesis that adult females not only represent the relational and decisional nucleus of the bonobo society, but also that they play a key role in affecting the emotional states of others.     

Social Attention in the Two Species of Pan: Bonobos Make More Eye Contact than Chimpanzees

Humans’ two closest primate living relatives, bonobos and chimpanzees, differ behaviorally, cognitively, and emotionally in several ways despite their general similarities. While bonobos show more affiliative behaviors towards conspecifics, chimpanzees display more overt and severe aggression against conspecifics. From a cognitive standpoint, bonobos perform better in social coordination, gaze-following and food-related cooperation, while chimpanzees excel in tasks requiring extractive foraging skills. We hypothesized that attention and motivation play an important role in shaping the species differences in behavior, cognition, and emotion. Thus, we predicted that bonobos would pay more attention to the other individuals’ face and eyes, as those are related to social affiliation and social coordination, while chimpanzees would pay more attention to the action target objects, as they are related to foraging. Using eye-tracking we examined the bonobos’ and chimpanzees’ spontaneous scanning of pictures that included eyes, mouth, face, genitals, and action target objects of conspecifics. Although bonobos and chimpanzees viewed those elements overall similarly, bonobos viewed the face and eyes longer than chimpanzees, whereas chimpanzees viewed the other elements, the mouth, action target objects and genitals, longer than bonobos. In a discriminant analysis, the individual variation in viewing patterns robustly predicted the species of individuals, thus clearly demonstrating species-specific viewing patterns. We suggest that such attentional and motivational differences between bonobos and chimpanzees could have partly contributed to shaping the species-specific behaviors, cognition, and emotion of these species, even in a relatively short period of evolutionary time.     

Co–Residence between Males and Their Mothers and Grandmothers Is More Frequent in Bonobos Than Chimpanzees

In long–lived social mammals such as primates, individuals can benefit from social bonds with close kin, including their mothers. In the patrilocal chimpanzee (Pan troglodytes spp.) and bonobo (Pan paniscus), sexually mature males reside and reproduce in their natal groups and can retain post-dependency bonds with their mothers, while immatures of both sexes might also have their paternal grandmothers available. However, quantitative information on the proportion of males and immatures that co-reside with both types of these close female relatives is limited for both species. Combining genetic parentage determination and group composition data from five communities of wild chimpanzees and three communities of wild bonobos, we estimated the frequency of co-residence between (1) mature males and their mothers, and (2) immature males and females and their paternal grandmothers. We found that adult males resided twice as frequently with their mothers in bonobos than in chimpanzees, and that immature bonobos were three times more likely to possess a living paternal grandmother than were immature chimpanzees. Patterns of female and male survivorship from studbook records of captive individuals of both species suggest that mature bonobo females survive longer than their chimpanzee counterparts, possibly contributing to the differences observed in mother–son and grandmother–immature co-residency levels. Taking into account reports of bonobo mothers supporting their sons'' mating efforts and females sharing food with immatures other than their own offspring, our findings suggest that life history traits may facilitate maternal and grandmaternal support more in bonobos than in chimpanzees.     

Bonobo but not chimpanzee infants use socio-sexual contact with peers

Bonobos have been observed to use socio-sexual behavior at higher frequency than chimpanzees. Little is known about the developmental influences that shape this behavior in bonobos. We compared the social sexual behavior of wild-born bonobo (n = 8) and chimpanzee (n = 16) infants in an experimental feeding test. Subjects of both species were orphans of the bushmeat trade living at sanctuaries in peer groups. During the experiment, chimpanzee infants never had socio-sexual interactions with one another. In contrast, bonobo infants had socio-sexual interactions significantly more than the chimpanzee infants and more often when food was presented. During these socio-sexual interactions, bonobo infants did not show a preference for heterosexual partners or genital–genital positioning that is reproductive in adults (e.g. a dorso–ventral posture). These findings suggest that the socio-sexual behavior previously observed in various captive and wild bonobos is species-typical. Wild-born bonobos originating from a large geographical range develop this behavior long before puberty and without the need for adults initiating such behavior or acting as models for observational learning. Meanwhile, chimpanzee infants of the same age with similar rearing history show no signs of the same socio-sexual behavior. Results are interpreted regarding hypotheses for the evolution of bonobo psychology.     

Reconciliation and Consolation in Captive Western Gorillas

We studied post-aggression mechanisms in a captive group of western gorillas (Apenheul Primate Park, The Netherlands) and compared them with those of wild mountain gorillas (Gorilla beringei). We found the same trend for reconciliation that wild mountain gorillas show: reconciliation occurred only between adult male-female dyads, while it was absent in the other sex-age class combination. There were both solicited and nonsolicited contacts; the latter finding is in contrast with the result obtained in wild mountain gorillas, in which consolation was absent. Immature females were more likely to offer consolation toward both related and unrelated individuals. Consolation did not reduce the likelihood of further attacks among group members. It may be that, as the α-male plays a fundamental role in preventing the spread of conflicts throughout the entire group, triadic contacts become ineffective for the function. The levels of consolation were higher in absence of reconciliation than in its presence, suggesting that consolation may function as an alternative mechanism in stress reduction of the victim.