Long‐term monitoring of social behavior in a grouping of six female tigers (Panthera tigris)

Although literature on the reproduction of captive tigers is plentiful, there is little information on other social behaviors, particularly within large social groupings. Here we report on the long‐term monitoring of social behaviors in six female tigers (Panthera tigris). Over a period of 6 years, behavioral data were collected on spraying, vocalizations, non‐contact aggression, and contact aggression during outdoor observations, as well as nearest neighbors in the morning, as a proxy measure of affiliation. Regression analyses showed that non‐contact aggression and vocalizing were increasing, whereas spraying and social proximity were decreasing over the 6 years of the study. Paired t‐tests showed no seasonal differences in aggression, but animals had higher social proximity scores during the mornings of the colder winter months. Correlations showed that spraying was positively related to contact aggression whereas social proximity was negatively correlated with non‐contact aggression. In addition to documenting levels of contact and non‐contact aggression over time, this study showed these behaviors correlated with behaviors that were more easily scored (spraying and social proximity). Behavioral monitoring has played a key role in the management and husbandry of this large social grouping of female tigers. In addition to providing keepers with the opportunity to observe their animals in regularly scheduled sessions, behavioral monitoring has provided baseline information on social relationships in this grouping over time. If future research validates the relationship between aggression and spraying/social proximity, this can be a valuable tool for long‐term monitoring of groups of tigers. Zoo Biol 27:89–99, 2008. © 2007 Wiley‐Liss, Inc.     

Chimpanzees Cope with Temporary Reduction of Escape Opportunities

Many studies have focused on the responses of primates to changes in population density, but little is known about the responses to more subtle changes in the physical environment. Studies on primate responses to high density supported the use of various coping strategies depending on the circumstances. We investigated which strategy was adopted by a large group of captive chimpanzees in response to a special case of space restriction: the temporary reduction of escape opportunities. Due to work on the outdoor enclosure, the chimpanzees at Chester Zoological Gardens were confined to their indoor quarters for 1 mo. As it was winter, the chimpanzees spent most of their time indoors also during the 7-week control period, but had the option to go outside. Therefore, there was no major difference in the actual density between the two conditions, but by being confined indoors the chimpanzees had reduced escape opportunities. Under this condition they would be expected not to alter their behavior dramatically but to adjust it in ways to avoid conflict and to reduce tension. The chimpanzees did not change their overall aggression rates during the period with a reduction of escape opportunities versus the control period. The patterns of allogrooming and submissive greetings also did not differ between the two periods. However, they appeared to refrain from joining ongoing aggressive interactions during the period with reduced escape opportunities, whereas no change in counteraggression occurred between the two periods. Our findings suggest that the chimpanzees may adopt a selective inhibition strategy when escape opportunities are limited. Further support for such selectivity comes from the analysis of different types of aggressor-target dyads. During the period with reduced escape opportunities, aggression rates decreased in dyads characterized by high aggression rates at baseline, i.e., high aggressor-target dyads, whereas the same individuals did not change their aggressive tendencies in low aggressor-target dyads. Ours is among the few studies showing a decrease, albeit selective, in aggressive behavior during a situation of space restriction. The ability to rapidly adopt an appropriate strategy and to inhibit aggression during such restriction seems to confirm findings of chimpanzees under high density conditions. The inhibition of aggressive tendency may be especially developed in chimpanzees and may be related to the natural challenges posed by the routine changes in subgroup membership characteristic of their fission-fusion social organization.     

The effects of temporary cover on the behavior of socially housed stumptailed macaques (Macaca arctoides)

To examine the effects of the provision of temporary cover on the behavior of stumptailed macaques (Macaca arctoides), a single group of 26 animals was observed during 25 testing periods over 7.5 months. During each testing period, the group was observed under two conditions. In the Cover condition, two solid temporary walls 9.6 m in length, were erected within the animals' living compound. In the No-Cover condition, the wall materials were stacked against one solid exterior wall of the compound. The provision of cover significantly reduced the levels of contact aggression, proximity between animals, and locomotion, and reduced the ability of the dominant male to monopolize copulations. It did not effect other measures of affiliation. Inconsistencies between these results and those of other published studies suggest that, while cover can have a significant influence on some kinds of social behavior in some situations, generalizations about the benefits of providing cover to captive animals may be premature.     

Conflict management by hamadryas baboons (Papio hamadryas hamadryas) during crowding: a tension-reduction strategy

Primates change a variety of behavioral responses during short-term exposure to crowding. Under crowded conditions, rates of aggression, submissive behavior, and affiliative behavior may increase or decrease. Different patterns of change among these three categories of response have been interpreted as various coping "strategies" for managing the increased risk of conflict under crowded conditions. Grooming is of particular interest because this behavior is known to have a calming influence on the recipient and could be used to manage conflict under tense situations. A captive group of nine hamadryas baboons (Papio hamadryas hamadryas) composed of two harems was observed under short-term crowding to determine whether this species adopts conflict-management strategies similar to those described in other primates. The aggression, submission, affiliation, and displacement activities of the six adults in the group (two males and four females) were recorded, and behaviors in their small indoor quarters was compared with baseline behaviors in the outdoor section of their enclosure, which had over 10 times more space. Repeated-measures analysis of variance (ANOVA) was used to compare behavioral rates during crowding with baseline rates recorded immediately after crowding and during matched controls collected at the same time of day as the crowding sessions. Aggression and submission rates did not change significantly across conditions. Huddling together and proximity increased during crowding, and females increased grooming of their harem male during crowding. Displacement activities (e.g., pacing and self-grooming) increased during crowding, but scratching, an indicator of anxiety in primates, did not. The pattern of behavior exhibited by this group conforms to an active "tension-reduction" strategy in which animals successfully reduce the higher risk of aggression during crowding.     

Species-specific patterns of nonapeptide brain gene expression relative to pair-bonding behavior in grouping and non-grouping cichlids

Comparative studies have revealed that vasopressin–oxytocin pathways are associated with both pair bonding and grouping behavior. However, the relationship between pair bonding and grouping behavior remains unclear. In this study, our aim was to identify whether two species that differ in grouping behavior display a corresponding difference in their pair bonds, and in the underlying vasopressin–oxytocin hormonal pathways. Using two species of cichlid fishes, the highly social Neolamprologus pulcher and the non-social Telmatochromis temporalis, we measured proximity of pairs during pair bond formation, and then measured social behaviors (proximity, aggression, submission, affiliation) and brain gene expression of isotocin and arginine vasotocin (the teleost homologues of oxytocin and vasopressin, respectively), as well as their receptors, after a temporary separation and subsequent reunion of the bonded pairs. Pairs of the social species spent more time in close proximity relative to the non-social species. Rates of aggression increased in both species following the separation and reunion treatment, relative to controls that were not separated. Overall, whole brain expression of isotocin was higher in the social species relative to the non-social species, and correlated with proximity, submission, and affiliation, but only in the social species. Our results suggest that both a social and a non-social cichlid species have similar behavioral responses to a temporary separation from a mate, and we found no difference in the brain gene expression of measured hormones and receptors based on our separation–reunion treatment. However, our results highlight the importance of isotocin in mediating submissive and affiliative behaviors in cichlid fishes, and demonstrate that isotocin has species-specific correlations with socially relevant behaviors.     

Coping with Acute Crowding by Cebus apella

Traditionally, scientists believed that crowded housing conditions increased aggression in animals. Recent research on captive primates fails to support a linear relationship between population density and aggressive behavior. Our study is the first to investigate the effects of acute crowding on a New World primate. We observed brown capuchins (Cebus apella) under 2 different spatial conditions: a control that offered the full indoor space in the enclosure and a short-term experimental condition that restricted the capuchins to approximately half the amount of space of the control condition. We compared rates of self-grooming, aggression, contact sitting, social grooming, and play between both conditions. We conducted a mixed factorial ANOVA at the individual level while taking subject sex and age class into account. Intense aggression, play, and social grooming decreased significantly in crowded conditions, suggesting that capuchins avoid social encounters if spatially confined. Their strategy for coping with acute crowding via a decrease in all forms of social behavior is intermediate between chimpanzee and macaque strategies in similar experiments. The only behavior to increase under crowding was a nonsocial, self-directed behavior with the potential of reducing arousal: self-grooming.     

Hormonal regulation of agonistic and affiliative behavior in female mongolian gerbils (Meriones unguiculatus)

Ovarian steroids and oxytocin (OT) have been implicated in the regulation of social behaviors. The purpose of the present study was to examine hormonal substrates of aggression and affiliation in the female Mongolian gerbil (Meriones unguiculatus), a highly social, monogamous rodent. Sexually naive adult females were paired with sexually experienced males for 48 h and their interactions videotaped. Females were gonadally intact and tested during vaginal estrus (INT) or ovariectomized and observed after the following treatments, administered by means of sc injections: EBEB (7 days of estradiol-benzoate); EBP (2 days of EB followed by progesterone), SALEB (saline, days 1-5 then 2 days of EB), OTEB (OT for days 1-5 then 2 days of EB); OTOIL (OT for days 1-5 then 2 days of OIL); or SALOIL (saline days 1-5 then 2 days of OIL). During the first hour of pairing INT females displayed higher levels of affiliation and lower levels of sniffing and agonistic behavior than SALOIL females. All hormonal treatments reduced agonistic behaviors when compared to SALOIL, although none of the hormonal treatments restored affiliation to INT levels. During the 48-h test overt aggression varied by treatment with INT, EBEB, EBP, and OTEB females displaying lower levels than SALOIL, while all groups displayed similar levels of affiliation. The results indicate that OT and E play a significant role in regulating male-directed aggressive behavior in females and that the presence of ovarian hormones as well as OT can increase affiliation during initial contact. Over a sustained period of cohabitation social cues appear to be more important in regulating affiliation than gonadal hormones.     

Dyadic and triadic reconciliation in pigtail macaques (Macaca nemestrina)

The tendency in primates for former antagonists to approach and affiliate following aggression has been termed reconciliation because the response is thought to resolve social conflicts produced by aggression. In primate societies, however, an aggressive interaction between two individuals often spreads to include other group members, especially the kin of the combatants. If post conflict affiliation resolves aggressive conflicts in a group, then affiliative increases might occur between combatants and the kin of their opponents following aggression as well as between former opponents. This hypothesis was tested in a captive group of 39 pigtail macaques (Macaca nemestrina) by comparing affiliative response frequencies of combatants during the 5 minute period following aggression to affiliative response frequencies during 5 minute baseline periods not preceded by aggressive activity. Following aggression, affiliation rates increased between combatants and their opponents, aggressors and the kin of their opponents, and aggressors and their own kin. Additional analyses showed that aggression among kin was reconciled more often than aggression among nonkin. Recipients of aggression reconciled with their attackers more often than aggressors reconciled with their victims. Animals with similar dominance ranks reconciled proportionately more often than those with large rank disparities and aggressive infractions of a calculated dominance hierarchy were reconciled more often than attacks consistent with the hierarchy. Results suggest that both dyadic and triadic reconciliations occur in M. nemestrina and that compared to other primate species M. nemestrina exhibit a moderate-to-high conciliatory tendency.     

Assessing the effects of social environment on blood pressure and heart rates of baboons

The objective of this publication is to report on the feasibility of using a tether system for obtaining data on blood pressure and heart rates of socially housed primates and to evaluate the extent to which housing environment alters cardiovascular responses (mean arterial blood pressure and heart rate). Blood pressure and heart rates of adult male baboons (Papio cynocephalus hamadryas) were evaluated over a 6 week period under three different housing conditions: social companion, individual, and socially unfamiliar. Social environment was manipulated in a specially designed cage that incorporated removable panels of either woven wire or solid sheet metal. The design of the cage permitted nonhuman primates to engage in species-typical social behaviors such as grooming and aggression. Using a tether and catheter system, we monitored cardiovascular physiology. We tested the hypothesis that individual housing, housing with social companions, and housing with social strangers would produce different mean arterial blood pressure and heart rate responses. Individual housing and housing with strangers produced resting mean arterial blood pressures that were elevated relative to blood pressure responses with social companions. Individual housing and housing with social strangers produced different patterns of cardiovascular response. Individual housing resulted in lowered heart rates and elevated blood pressures relative to the social companion condition. Housing with social strangers resulted in both elevated blood pressure and elevated heart rate, relative to the social companion condition. Responses observed during this study demonstrated the sensitivity of blood pressure and heart rates to differences in social environment.     

Environmental effects on the behavior of zoo-housed lions and tigers, with a case study of the effects of a visual barrier on pacing

Tigers and lions in the wild are nocturnal nonhuman animals who may hunt and mate opportunistically during daylight hours. In captivity, they spend most time on exhibit sleeping or pacing. To better understand their activity budget, this study examined the daily behavior patterns of 2 Sumatran tigers and 3 African lions in different housings. The proportion of scans the large felids spent engaged in stereotypic pacing varied by time of day and environment. The tigers spent different amounts of time pacing when housed in different exhibits; the lions paced more in off-exhibit housing than when on exhibit. These differences suggest changes to the cats' immediate housing environment may decrease pacing but provide little insight into altering specifics. Carnivores' pacing relates to their inability to control sensory access to social partners. Both environments with increased pacing contained chain-link fencing. allowing uncontrolled sensory contact. Where the tigers paced, the study placed a visual barrier between one female and keepers' or conspecifics' cues. This did not significantly decrease pacing. However, the study suggests considering sensory access and environmental variables when designing environments for captive carnivores.     

Environmental Effects on the Behavior of Zoo-housed Lions and Tigers,with a Case Study o the Effects of a Visual Barrier on Pacing

Tigers and lions in the wild are nocturnal nonhuman animals who may hunt and mate opportunistically during daylight hours. In captivity, they spend most time on exhibit sleeping or pacing. To better understand their activity budget, this study examined the daily behavior patterns of 2 Sumatran tigers and 3 African lions in different housings. The proportion of scans the large felids spent engaged in stereotypic pacing varied by time of day and environment. The tigers spent different amounts of time pacing when housed in different exhibits; the lions paced more in off-exhibit housing than when on exhibit. These differences suggest changes to the cats' immediate housing environment may decrease pacing but provide little insight into altering specifics. Carnivores' pacing relates to their inability to control sensory access to social partners. Both environments with increased pacing contained chain-link fencing. allowing uncontrolled sensory contact. Where the tigers paced, the study placed a visual barrier between one female and keepers' or conspecifics' cues. This did not significantly decrease pacing. However, the study suggests considering sensory access and environmental variables when designing environments for captive carnivores.     

Postconflict third-party affiliation in rooks, Corvus frugilegus

Conflict features in the lives of many animal species and induces social stress mediated by glucocorticoid hormones [1]. Postconflict affiliation, between former opponents (reconciliation) or between former opponents and a bystander (third-party affiliation), has been suggested as a behavioral mechanism for reducing such stress [2], but has been studied almost exclusively in primates [3]. As with many primates, several bird species live in social groups and form affiliative relationships [4]. Do these distantly related animals also use affiliative behavior to offset the costs of conflict? We studied postconflict affiliation in a captive group of rooks. Unlike polygamous primates, monogamous rooks did not reconcile with former opponents. However, we found clear evidence of third-party affiliation after conflicts. Both initiators and targets of aggression engaged in third-party affiliation with a social partner and employed a specific behavior, bill twining, during the postconflict period. Both former aggressors and uninvolved third parties initiated affiliative contacts. Despite the long history of evolutionary divergence, the pattern of third-party affiliation in rooks is strikingly similar to that observed in tolerant primate species. Furthermore, the absence of reconciliation in rooks makes sense in light of the species differences in social systems.     

Postweaning environmental and social factors influencing the onset and expression of agonistic behavior in Norway rats

Two studies were conducted to determine the importance of the postweaning environment and social milieu in regulating the expression of intraspecific aggression in Norway rats. In Experiment 1, male rats were housed either individually or in pairs at 21 days of age. In addition, one-half of the singly housed and paired animals were given experiences with intruders during maturation. At 85 days of age, all animals were given a brief intruder test and then removed from their postweaning environment and provided individually with homecages for a two week period until tested for aggression toward intruders. Results of intruder tests given during maturation indicated agonistic exchanges appeared earlier and more frequently in cages housing a single resident than cages with cohabiting males. However, agonistic exchanges between singly reared residents and intruders had detrimental consequences in adulthood especially under long-term combat situations. That is, although individually reared animals, with early fighting experiences, were capable of initiating intraspecific aggression, such individuals were unable to defend their homecage over a long period of time as evidenced by the high number of wounds and tendency to lose body weight during adult fighting.In Experiment 2, male Norway rats were reared in pairs from 21 days of age and identified as dominant or subordinate on the basis of intracolony social interactions shown during maturation. At 80 days of age, animals were paired with individually reared males in an unfamiliar cage for a 20 day period and examined for agonistic behavior toward intruders at 100 days of age. Group-reared subordinate males exhibited defensive behavior during confrontations with individually reared animals and incurred more wounds and lost more body weight than their cohabiting partner. In addition, subordinate males showed significantly fewer offensive postures toward intruders than individually reared cohabitants. In contrast, group-reared dominant animals did not differ from individually reared males in display of agonistic patterns, in number of wounds, and body weight changes during the period of cohabitation. These findings demonstrate that early rearing factors have pronounced effects on agonistic behavior. Animals experiencing defeat during development are more likely to lose agonistic confrontations in unfamiliar territory than either animals dominant in their early social interactions or animals without the experience of winning or losing agonistic encounters. These results have implications for the understanding of agonistic behavior and predicting outcomes of animal contests, and reveal important differences in agonistic experiences among animals reared in groups.     

The Long-Term Effects of Reconciliation in Japanese Macaques Macaca fuscata

With one exception, all previous studies of reconciliation in non-human primates (friendly reunion between former opponents) have focused on demonstrating the immediate, short-term effects despite the widely held view that reconciliation has a long-term function of repairing social relationships following aggression. To investigate this long-term function I compared mean interaction rates between opponents during the 10 d following reconciled and non-reconciled conflicts to baseline levels of interaction. Aggression rates during the 10 d after non-reconciled conflicts were significantly higher than the baseline rate, whereas after reconciled conflicts aggression was minimal. Similarly, grooming, proximity and approach rates during the 10 d after non-reconciled conflicts were significantly lower than the baseline rate whereas grooming, proximity and approach rates in the 10 d after reconciled conflicts were restored to baseline levels. These results indicate that there are consequences to not reconciling with a former opponent and highlight the fact that these may be costly in terms of increased risk of long-term aggression and reduced affiliation. The data support predictions from the Relationship-Repair Hypothesis suggesting that reconciliation functions as a mechanism for the repair of social relationships damaged by aggression.     

Social Relationships in Free-Ranging Male Macaca arctoides

Macaque social relationships differ greatly between species. Based on captive studies that focus mainly on females, researchers have classified stumptail macaque (Macaca arctoides) social relationships as tolerant, as indicated by a high rate of affiliation, frequent aggression, and symmetrical conflicts. To accumulate more data on male social relationships, which are relatively understudied, and to generate comparative data, we investigated male social relationships in a provisioned group of 68 free-ranging, naturally dispersing stumptail macaques in southern Thailand. We collected continuous focal animal and ad libitum data on 7 adult and 2 subadult males, recording social behavior during 283 contact hours between December 2006 and March 2007. Stumptail macaques of this population were less tolerant than predicted based on previous studies on captive groups: Rates of spatial proximity, affiliation, and aggression were low, most males directed affiliative behavior toward higher-ranking males, and conflicts were generally of low intensity and relatively asymmetrical. Thus, male stumptail macaques of the focal group appear to differ in their social style from a previous study of a captive group that mainly comprised of females. In some traits, they are even more intolerant than rhesus macaques, an intensively studied intolerant macaque species. We also compare our data on stumptail macaque males to those on other male macaques, but available data are too sparse to draw final conclusions.     

Rook,But Not Jackdaw,Post‐Conflict Third‐Party Affiliation Reduces Aggression for Aggressors

Post‐conflict (PC) affiliation refers to positive social interactions that occur after fights. Although this behavior has been widely studied, its functions are rarely tested. We examine a potential function of PC third‐party affiliation (affiliation between former opponents and bystanders) in rooks and jackdaws by investigating the hypothesis that conflicts lead to further aggression and that PC third‐party affiliation increases to reduce such aggression. The results show that PC affiliation reduces PC aggression for rook aggressors who were less likely to receive aggression after conflicts when they were affiliating with another vs. when they were alone. The opposite result was found for victims of both species who received more aggression after conflicts, and this aggression was not reduced by the act of affiliating. Finally, for jackdaw aggressors, the amount of aggression received after conflicts was not influenced by whether the individual was affiliating or alone, indicating that PC third‐party affiliation may serve a function that we did not examine. These findings highlight the importance of investigating functional differences in PC affiliative behavior according to the role played in the conflict.     

Experimental Manipulation of Food Accessibility Affects Conflict Management Behaviour in Ravens

Conflict management strategies such as reconciliation and bystander affiliation have been described for a variety of species. A common determinant seems to be a ‘complex’ social life, with individuals relying on affiliate relationships or social bonds. Little is known, however, about the strategic and flexible use of conflict management skills in experimental settings in species other than primates. We here investigated conflict and post‐conflict behaviour of ravens by manipulating the accessibility of food and, thus, the likelihood of aggressive interactions while foraging. Specifically, we presented birds with a certain amount of highly preferred food that varied in the number of pieces (one piece, two pieces or, as a control, small pieces matching the number of participating birds) and observed their agonistic behaviour during feeding and their affiliative behaviour afterwards. The results showed that high levels of conflicts during feeding in the 1‐piece and 2‐piece conditions led to high levels of affiliation after feeding. Depending on the experimental condition, this effect is best explained (a) by the affiliative behaviour of former aggressors (1‐piece condition) and (b) by the affiliation directed to the receivers of aggression after feeding (2‐piece condition). Those dyads that engaged in allo‐preening after feeding also engaged in allo‐preening outside the experimental setting, suggesting that socially bonded individuals provided third‐party affiliation to victims of aggression. Moreover, socially bonded ravens fed close to each other in the experiment when food was clumped, indicating that they actively coordinated their behaviour when there was a high conflict potential. Taken together, these findings support the assumption that ravens use their social bonds to avoid conflicts by choosing with whom to feed, and to buffer effects of conflicts by engaging in third‐party affiliation as post‐conflict behaviour.     

The effects of transport stress on tiger physiology and behavior

Tigers are often transported for education, conservation, and zoo enhancement purposes, however the effect of transfer on them has not yet been documented. Our objective was to evaluate how transport affects the behavior and physiology of tigers, taking into account previous experience with the transport procedure. We simulated transport by relocating five tigers in a small individual transfer cage. Two tigers had prior experience with the procedure, and three tigers were naïve to it. After 30 min, each tiger was released back into their original enclosure. Physiological measurements were recorded for four of the five tigers; these included respiration rate and immune‐reactive fecal cortisol response using radioimmunoassay. We also recorded the behavior of all five tigers before, during, and after transport. Our behavioral analysis included activity level, pacing behavior, time spent investigating, respiration rate, and ear position. Average respiration rates of all tigers increased from 56.1 breaths/min to 94.6 breaths/min during transport and to 132.3 breaths/min 10 min following release into their enclosures. Average immune‐reactive cortisol concentrations peaked 3–6 days after transport at 239% above baseline and returned to baseline levels 9–12 days afterward. During their peak time block, naïve tigers exhibited a higher average increase in cortisol levels (482% above baseline) than the experienced tigers (158% above baseline). The naïve tigers' average immune‐reactive cortisol concentration remained elevated for a longer period (9–12 days) than the experienced tigers' (3–6 days). In both groups, behavioral responses ranged from active to inactive, however naïve tigers performed these repertoires with greater intensity by pacing faster and performing fewer state changes. Results suggest that prior exposure to elements of the transport procedure may lead to some level of habituation, thus reducing the effects of transportation stress. Zoo Biol 23:335–346, 2004. © 2004 Wiley‐Liss, Inc.     

Post-conflict affiliation by chimpanzees with aggressors: other-oriented versus selfish political strategy

Consolation, i.e., post-conflict affiliation directed from bystanders to recent victims of aggression, has recently acquired an important role in the debate about empathy in great apes. Although similar contacts have been also described for aggressors, i.e., appeasement, they have received far less attention and their function and underlying mechanisms remain largely unknown. An exceptionally large database of spontaneous conflict and post-conflict interactions in two outdoor-housed groups of chimpanzees lends support to the notion that affiliation toward aggressors reduces the latter's aggressive tendencies in that further aggression was less frequent after the occurrence of the affiliation. However, bystander affiliation toward aggressors occurred disproportionally between individuals that were socially close (i.e., affiliation partners) which suggest that it did not function to protect the actor itself against redirected aggression. Contrary to consolation behavior, it was provided most often by adult males and directed toward high ranking males, whereas females engaged less often in this behavior both as actors and recipients, suggesting that affiliation with aggressors is unlikely to be a reaction to the distress of others. We propose that bystander affiliation toward aggressors may function to strengthen bonds between valuable partners, probably as part of political strategies. Our findings also suggest that this post-conflict behavior may act as an alternative to reconciliation, i.e., post-conflict affiliation between opponents, in that it is more common when opponents fail to reconcile.     

Golden-bellied mangabeys (Cercocebus galeritus chrysogaster) at the Sacramento Zoo: Adult heterosexual pairs

The behaviors of three different adult male–adult female pairs of golden-bellied mangabeys (Cercocebus galeritus chrysogaster) were compared. Correlates of restricted social conditions and of the presence of offspring on adult behavior and direction of aggressive behaviors were noted. A captive pair displayed self-biting and other aberrant behaviors, a lone wild-caught pair showed redirected aggression related to sex, and a pair housed with its offspring showed close social proximity and protection of an infant. The implications of housing and experience for space and visitor exposure are discussed. Observer- vs. visitor-directed displays are seen as important measures of group and visitor effects.