Nonaggressive interventions by third parties in conflicts among captive Bornean orangutans (<Emphasis Type="Italic">Pongo pygmaeus</Emphasis>)

Whereas orangutans are regarded as semisolitary animals in the wild, several studies have reported frequent social interactions, including aggression, among orangutans in captivity. As yet, there is a lack of knowledge about how they cope with aggression. In this report, we provide a number of new observations of interventions by third parties in aggressive interactions within a captive group of Bornean orangutans (Pongo pygmaeus) in the Tama Zoological Park, Japan. We observed that an adult female and a juvenile male orangutan intervened in aggressive interactions. The victim was a newly introduced juvenile female who was unrelated to anyone in the zoo. The ways in which the orangutans intervened were not aggressive, as the interveners simply aimed to separate the opponents, and these interventions did not lead to further aggression in almost every case. Our observations suggest that third parties can play an important role in managing aggressive conflicts among captive orangutans and, under conditions in which orangutans share limited space, nonaggressive interventions by third parties for settling conflicts appear. It is possible that orangutans may actively promote the peaceful coexistence of other individuals.     

Reconciliation in captive cotton‐top tamarins (Saguinus oedipus), a cooperative breeding primate

Reconciliation has been demonstrated in all primate species in which the phenomenon has been studied. However, reconciliation has been studied in only two species of callitrichids, and conclusions remain controversial. The first aim of this study has been to find out whether captive cotton‐top tamarins (Saguinus oedipus) reconcile, since this is the first such study on this species. We examined 227 conflicts in three family groups (N=19). Instances in which individuals remained together in t=0 (29; 12.8%) were not analyzed. The cotton‐top tamarins showed heightened affiliation between opponents in the postconflict periods (PC) compared with matched control (MC) periods (39.88±5.12% and 3.18±1.27%, respectively), with a corrected conciliatory tendency of 37.17±5.37%, and a “time window” that included the first 180 sec of the PC period. Former opponents were the most likely recipient of affiliative behaviors during the PC periods: 39.83±4.26% vs. 11.36±5.33% during MC periods. The proportion of attracted pairs (47.13±6.25%) was significantly higher than those of dispersed pairs for male–male conflicts (3.79±1.79), but not for male–female conflicts (27.31±9.32 and 4.82±2.9, respectively). In cooperative‐breeding species, specific sex‐class dyads might differ in how they resolve conflicts. Am. J. Primatol. 71:895–900, 2009. © 2009 Wiley‐Liss, Inc.     

Reconciliation in wild white-faced capuchins (Cebus capucinus)

The likelihood of reconciliation (defined as preferential peaceful contact among former opponents following conflicts) has been predicted to vary positively with relationship value and compatibility, and negatively with relationship security. Long-term data on wild white-faced capuchins (Cebus capucinus) indicate that dyads consisting of an adult female and an alpha male have high value and compatibility, but low security. Two studies of C. capucinus postconflict (PC) behavior were carried out at Lomas Barbudal Biological Reserve, Costa Rica. One study consisted of 30-min PC and matched control (MC) follows. The second study extracted PC and MC periods from long follows, yielding PC/MC periods averaging 105 min. In study 2, but not study 1, significantly more PC/MC pairs were attracted (former opponents affiliated with each other sooner in the PC period than in the MC period) than were dispersed (former opponents affiliated with each other sooner in the MC period than in the PC period). Reconciliation in study 2 could not be explained as a by-product of former opponents' tendency to seek affiliative contact with conspecifics generally, or of the spatial proximity of opponents following conflicts. Attempted reconciliation was less likely to be followed by renewed aggression when reconciliation attempts were delayed following conflicts. The data were insufficient for a formal test of differences in conciliatory tendency (the difference between the number of attracted and dispersed PC/MC pairs, divided by the total number of pairs) among dyad types to be conducted.     

Social influences on first reproductive success and related behaviors in the saddle-back tamarin (Saguinus fuscicollis,callitrichidae)

Some aspects of sociosexual behavior and the age at which maturing females experienced their first evident pregnancy and at which maturing males caused their first evident pregnancy were recorded in Saguinus fuscicolliscohabiting from 6 months of age with either an adult or a maturing sex partner. The following pair combinations and trios were studied: young male -young female, young male-adult female, adult male-young female, and adult male-young male-young female. The most frequent type of social interaction between young animals was rough and tumble play, while huddling was the most frequent interaction between young animals and their adult partners. Grooming and sexual interactions were very infrequent and there were no differences in the frequencies of these interactions among subject groups. Maturing females cohabiting with an adult male conceived significantly earlier than maturing females cohabiting with a male of their own age. Maturing males cohabiting with adult females sired offspring at a significantly earlier age than males cohabiting with a female of their own age. Some possible behavioral and physiological processes involved in the causation of early reproductive success in young tamarins cohabiting with adults are discussed. Saguinus fuscicollis fuscicollis andSaguinus fuscicollis illigeri.     

Conflict and postconflict behaviour in captive black-and-white snub-nosed monkeys (<Emphasis Type="Italic">Rhinopithecus bieti</Emphasis>)

Black-and-white snub-nosed monkeys (Rhinopithecus bieti) have almost never been the subject of any behavioural observations in captivity. This study was aimed at providing preliminary information about agonistic and reconciliation behaviour in a group kept at the Kunming Institute of Zoology in China. Established procedures were used for this investigation (i.e., the postconflict/matched-control method and the time-rule method). Intra-group aggression rates were quite low. Postconflict affiliation as well as selective attraction of former opponents to each other following conflicts was demonstrated. Former opponents contacted each other earlier in postconflict periods than in matched-control periods. The average conciliatory tendency of all focal individuals combined was 54.5%. After an agonistic interaction, the first affiliative contact between former aggressors usually took place within the first minute. The behaviours most often shown as first affiliations after a conflict were body contact, mount, touch, and hold-lumbar, of which the latter is an explicit reconciliatory gesture. Furthermore, the adult male intervened non-aggressively in 84% of all conflicts (n=25) among the adult females. Overall, the patterns of aggression and reconciliation observed in R. bieti bear many of the traits that characterise tolerant primate species.     

The effect of inter-opponent distance on the occurrence of reconciliation in stumptail (Macaca arctoides) and rhesus macaques (Macaca mulatta)

Several studies have shown that affiliative contact between opponents increases after conflicts in a number of primate species. This study investigated the effect of inter-opponent distance on the occurence of reconciliation in stumptail and rhesus macaques. Both species were represented by multi-male, multi-female groups housed in large outdoor compounds. Ten-minute focal animal observations during post-conflict (PC) and matched control (MC) samples were collected in which affiliative interactions and the inter-opponent distance at the beginning of each sample were recorded. A total of 251 and 561 PC-MC opponent pairs were obtained for stumptail and rhesus macaques, respectively. Inter-opponent distance was smaller in PC than in MC observations for both stumptail and rhesus macaques. Conciliatory tendency varied as a function of the initial PC-MC inter-opponent distance. In particular, the closer opponents were in PC compared to MC periods, the more likely they were to reconcile. When the inter-opponent distance was the same in PC and MC periods the conciliatory tendency of both species was notably reduced. These findings have implications for the estimation of conciliatory tendency and its use in comparative studies.     

Immature and adult langur monkey (Presbytis entellus) males: Infant-initiated adoption in a colony group

This paper focuses on interactions of an immature male langur monkey (Presbytis entellus) with an adult male in a captive social group. During his mother's 2-week absence, an 8-month-old male actively sought and received care from two adults, including an adult male. The interactions of the 2 males are compared with the behavior of 12 other adult male-immature male pairs to illustrate variability in male-male relations for a species in which adult male-young interactions are reported to be minimal and in which males normally spend portions of their lives in all-male groups.     

Sex of opponent influences response to a potential status signal in house sparrows

We evaluated the possible use of a sexually dimorphic plumage trait as a status signal in inter- and intrasexual social interactions in free-living house sparrows, Passer domesticus. Males with larger black throat patches (bibs) tended to be dominant regardless of the opponent's sex. This relationship was maintained after controlling for body size and age in multivariate analyses. However, the nature of social interactions varied according to the sex of the opponent and a male's bib size. Males with larger bibs tended to be less aggressive to male opponents, and received significantly less aggression from those birds, a key predicted benefit of a status signal. However, when the opponent was a female, males with large bibs displayed significantly more aggression than males with small bibs, and this relationship persisted even after controlling for the dominance status of the participants. Females were significantly more aggressive towards male opponents than towards female opponents, and females interacting with two males of known bib size were more aggressive to the one with the larger bib. These results suggest that the bib of male house sparrows may be a signal of status, but that in interactions with females, the bib may also have other functions. Frequent testing by females of the underlying quality being signalled by the bib may be linked to later mate choice, and might be adaptive if the information value of the bib varies among years. Such testing may also contribute to mechanisms for maintaining the bib as a reliable signal of status. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

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.     

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.     

Affiliative relationships between adult males and immature group members in naturally occurring ringtailed lemurs (Lemur catta)

Affiliative relations between adult males and immature group members were studied in three naturally occurring groups of ringtailed lemurs (Lemur catta) over a 12-month period at Beza-Mahafaly Reserve, southwestern Madagascar. No statistically significant difference was found in rates of affiliative interactions between adult males and older immatures (juveniles and adolescents) over reproductive seasons. However, some of the adult males in two focal groups increased their rates of affiliative behaviors with older immatures during the lactation and subsequent 1993 postmigration periods. Female involvement with infants during lactation season, the increasing independence of juveniles and adolescents, and length of male tenure may be factors contributing to such a pattern. Neither adult males nor immatures were significantly responsible for the initiation and maintenance of proximity in male-immature dyadic interactions. Neither dominance rank nor age-class of the adult male affected their rates of affiliative behavior with immatures. The majority of focal males exhibited an interest in infants and occasionally participated in alloparental care. It is suggested that adult males can benefit from affiliative relations with immatures in terms of opportunities for greater spatial centrality in the group, which can lead to enhanced predator protection and greater opportunities to develop affiliative relationships with females. Immatures can benefit from affiliation with males in relation to enhanced predator detection and protection, alloparental care, and opportunities to develop social skills. Am J Phys Anthropol 103:163–171, 1997. © 1997 Wiley-Liss, Inc.     

The effect of hunger on mating behaviour and sexual selection for male body size in Gerris buenoi

The set of mating behaviours expressed by an individual may depend upon the state of that individual and local environmental conditions. Understanding how these factors affect mating behaviours may elucidate how a mating system operates, and its consequences for the form and strength of sexual selection. We conducted two experiments on the water striderGerris buenoi to (1) determine the effect of hunger on the mating behaviour of both sexes and (2) examine female choice for large males. In our first experiment, we manipulated hunger (20 h starvation) in both sexes and recorded mating, male harassment, copulation duration and guarding duration. We predicted that hunger would increase female reluctance to mate because mating conflicts with foraging. Female hunger (20 h starvation) decreased mating rate by two-thirds but had no significant effect on male mating behaviour. In a second experiment, we examined the effect of female hunger, and resulting reluctance, on sexual selection for large male size. Hungry females (5 h starvation) were placed with two fed males (one large, one small) and we recorded male premating and mating behaviours. We observed significant large-male mating advantage when females were hungry, but not when satiated. Mating efforts (harassment, premating struggles) were similar for both male phenotypes in both female hunger treatments, suggesting that the mating advantage of large males resulted from increased reluctance of hungry females to mate. Neither male body size nor female hunger explained a significant amount of variation in copulation duration or guarding duration. We discuss our results in light of two competing hypotheses for female choice (active and passive) on male body size and suggest that passive choice for large males acts in this system. Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Post-conflict affiliation in common marmosets (Callithrix jacchus jacchus)

Post-conflict (PC) affiliation has been demonstrated in a number of Old World monkeys and apes, but very little is known about the occurrence of the phenomenon in New World monkeys. This study examined 282 PC interactions after spontaneous conflicts around feeding time in two family groups (N = 12) of captive common marmosets, Callithrix jacchus jacchus. We found an overall corrected conciliatory tendency of 31%. Selective attraction was seen: former opponents met significantly more often in PCs than in matched controls (MCs) (27% and 16%, respectively). There was no difference in the occurrence of PC affiliation between dyads consisting of parent-offspring constellations compared to offspring-offspring constellations. PC affiliative behaviors were seen in the first three minutes following conflict termination, and consisted mainly of proximity, play invitations, and food transfer. Notably, former opponents remained within arm's reach after 17% of conflicts. Affiliation was more likely to follow after conflicts involving play issues. The functional importance of the PC affiliation in marmosets remains to be examined.     

The impact of paternity on male–infant association in a primate with low paternity certainty

In multimale groups where females mate promiscuously, male–infant associations have rarely been studied. However, recent studies have shown that males selectively support their offspring during agonistic conflicts with other juveniles and that father's presence accelerates offspring maturation. Furthermore, it was shown that males invest in unrelated infants to enhance future mating success with the infant's mother. Hence, infant care might provide fitness gain for males. Here, we investigate male–infant associations in rhesus macaques (Macaca mulatta), a primate with low paternity certainty as females mate with multiple partners and males ensure paternity less efficiently through mate‐guarding. We combined behavioural data with genetic paternity analyses of one cohort of the semi‐free‐ranging population of Cayo Santiago (Puerto Rico) and recorded affiliative and aggressive interactions between focal subjects and adult males from birth to sexual maturation (0–4 years) of focal subjects. Our results revealed that 9.6% of all interactions of focal subjects involved an adult male and 94% of all male–infant interactions were affiliative, indicating the rareness of male–infant aggression. Second and most interestingly, sires were more likely to affiliate with their offspring than nonsires with unrelated infants. This preference was independent of mother's proximity and emphasized during early infancy. Male–infant affiliation rose with infant age and was pronounced between adult males and male rather than female focal subjects. Overall, our results suggest that male–infant affiliation is also an important component in structuring primate societies and affiliation directed towards own offspring presumably represent low‐cost paternal care.     

Coalition formation among male Barbary macaques (Macaca sylvanus)

A coalition is formed when one animal intervenes in an ongoing conflict between two parties to support one side. Since support of one party is also an act against the other party, coalitions are triadic interactions involving a supporter, a recipient, and a target. The purpose of this study was to test which of three possible theories explains coalition formation among male Barbary macaques: 1) Males support kin to enhance their indirect fitness (kin selection). 2) Males support nonkin to receive future reciprocal support (reciprocal altruism). 3) Males pursue self-interests and immediately benefit via nonkin support (cooperation). Coalition formation was investigated among 31 semi-free male Barbary macaques in the Salem Monkey Park, Germany during the mating season. The results show: 1) Males intervened more often in dyadic conflicts in which a related opponent was involved and supported related opponents more than unrelated opponents. Close kin supported each other more often than distant kin. 2) Some evidence for reciprocal support was found. However, reciprocity was probably a by-product of targeting the same individuals for dominance. 3) Coalition formation among nonkin is best interpreted as cooperation, based on self-interests. Male Barbary macaques seem to intervene more often to stabilize and less often to improve their rank. Although our data were limited, the results revealed that kin support, reciprocal support, and cooperative support were all involved in coalition formation among male Barbary macaques.     

Does Male Care,Provided to Immature Individuals,Influence Immature Fitness in Rhesus Macaques?

Among many mammals, maternal care strongly impacts infant survival; however, less is known about whether adult males also affect infant fitness. Paternal care is expected when providing care enhances offspring survival and reproduction, which likewise increases fathers’ fitness. Males might also care for unrelated immature individuals to increase their mating probability with the immature individuals’ mothers. Studies in multimale primate groups showed that sires enhance food access for offspring and provide protection in conflicts. Furthermore, fathers’ presence during infancy has been suggested to accelerate offspring sexual maturation. However, no study has yet directly linked the degree of father-offspring bonds to offspring fitness in primates. We previously reported father-offspring affiliation in rhesus macaques, pronounced during early infancy and independent of mothers’ presence. The present study aims at investigating whether affiliation with fathers or other males affects proxies of immature fitness (body mass gain, body fat and testis size). First, we combined behavioral, genetic and morphometric data from 55 subjects of one group. Second, using demographic and genetic data, we investigated for 92 individuals of the population whether mother- and father-offspring co-residence during immaturity influenced offspring lifetime reproductive success (LRS). Our results show that focal rank and higher amounts of affiliation with high-ranking males during infancy tend to positively impact body mass gain of female, but not male focal animals. In contrast, body mass gain of male focal individuals, but not females’, appeared to be higher when affiliation of male immature individuals was evenly distributed across their adult male partners. Moreover, we found mothers’, but not fathers’, presence during immaturity to predict offspring LRS. Our results suggest that male-immature affiliation, but not father-offspring co-residence, potentially impacts proxies of immature fitness. However, future studies should investigate the underlying mechanisms of male-immature relationships and their impact on immature fitness in more detail.     

Mycophagous beetle females do not behave competitively during intrasexual interactions in presence of a fungal resource

Intrasexual interactions can determine which individuals within a population have access to limited resources. Despite their potential importance on fitness generally and mating success especially, female–female interactions are not often measured in the same species where male–male interactions are well‐defined. In this study, we characterized female–female interactions in Bolitotherus cornutus, a mycophagous beetle species native to Northeastern North America. We used dyadic, behavioral assays to determine whether females perform directly aggressive or indirectly exclusionary competitive behaviors. Polypore shelf fungus, an important food and egg‐laying resource for B. cornutus females, is patchily distributed and of variable quality, so we tested for competition over fungus as a resource. Behavior of females was assessed in three sets of dyadic trials with randomly paired female partners. Overall, females did not behave aggressively toward their female partner or perform exclusionary behaviors over the fungal resource. None of the behaviors performed by females were individually repeatable. Two scenarios may explain our lack of observed competition: our trial context may not induce competition, or female B. cornutus simply may not behave competitively in the wild. We compare our results to a similar study on male–male interactions in the same species and propose future studies on female–female interactions under different competitive contexts to expand the understanding of female competition.     

Post-conflict Behaviour of Wild Olive Baboons. II. Stress and Self-directed Behaviour

Previous studies indicate that the proximate function of post-conflict affiliative interaction among primates is to reduce a victim's uncertainty about its opponent's future behaviour: the ‘uncertainty reduction hypothesis’ (Aureli & van Schaik 1991; Ethology, 89 , 101–114). This study confirms and extends these results, demonstrating that they are neither a product of captivity nor specific to macaques: both victims and initiators of aggression in a large group of wild olive baboons exhibited elevated rates of self-directed behaviour (SDB) - scratching, autogrooming, body-shaking and yawning - in a 10-min post-conflict period. During this period, they were more likely to receive further aggression. Reconciliation reduced both SDB and the incidence of further aggression. However, reconciliation only reduced SDB among individuals involved in conflicts in which both parties exchanged aggression. It is suggested that aggressors in unilateral conflicts were aroused rather than uncertain and that their victims' lack of control over post-conflict interactions (which tended to be initiated by their opponents) prevented them from benefiting from reconciliation in the same fashion as longtailed macaque victims, which frequently initiated reconciliation.     

Seasonal changes in and effects of familiarity on agonistic behaviors of rat-like hamsters (Cricetulus triton)

The seasonal changes in agonistic behaviors and effects of familiarity on agonistic behaviors in wild-caught adult rat-like hamsters (Cricetulus triton) were observed in dyadic encounters in a neutral arena. The aggression of opposite- and same-sex encounters became higher or remained the same during the non-breeding season. This indicates that the hamsters were solitary during both seasons. Familiarity increased the aggression in male–male encounters and decreased the aggression in female–female encounters during both seasons. Familiarity also increased the aggression in female–male encounters during the non-breeding season and had no effect on the aggression in female–male encounters during the breeding season. These results may be related to the hamsters social structure. The more agonistic acts both male and female hamsters had, the more frequently they marked using flank glands during both seasons. This implies that flank gland marking can be used to advertise status and can be assessed by opponents to reduce the agonistic costs.     

Carotenoid-based epaulettes reveal male competitive ability: experiments with resident and floater red-shouldered widowbirds

Many birds display carotenoid-based ornaments, which are typically considered to be honest indicators of individual health and condition. Experimental work on male red-shouldered widowbirds, Euplectes axillaris, has demonstrated a function of the carotenoid-based epaulettes in male contests and territory acquisition. Using two experiments, we investigated whether the natural variation in this colour signal reveals male competitive ability. Males with larger and redder (more longwave) epaulettes established territories to the exclusion of males with smaller and less red signals, which formed a large population of ‘floaters’. In an experiment in which we removed 42 resident males, these floaters rapidly filled up vacant territories. Among removed birds held in captivity, residents strongly dominated floaters in dyadic contests over access to an easily monopolized feeder (i.e. outside the context of territory defence). Only epaulette size predicted the outcome of these male contests. In addition, when competitors were experimentally given similar epaulette signals (removed or painted red to the average population size), the males were involved in more aggressive interactions than during unmanipulated contests, but residents continued to outcompete floaters. On release (after 8 days) to the breeding grounds, most residents (88%) rapidly reclaimed their territories from replacements. Combined, these results suggest that some intrinsic ‘resource-holding potential’, associated with the variation in epaulette signal, is primarily responsible for residents dominating nonresidents. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.