Aggression with mixed age-sex groups of bolivian squirrel monkeys following single animal introductions and new group formations

The social organization of the Bolivian squirrel monkey (Saimiri boliviensis boliviensis) is thought to be sexually segregated, with males and females forming separate social groups during the nonbreeding season. To investigate the influence of this social order on patterns of aggression, controlled single animal introductions within established groups and establishment of new groups were studied in a systematic manner. Behavioral observations were made using an all-occurrences technique to sample all agonistic interactions. In study I, 4 animals of each age-sex class (adult males and females, juvenile males and females) were introduced one at a time into 4 different social groups composed of 1 adult male and 8 to 10 adult females. Behavioral observations were made prior to and after the introductions. Newly introduced adult males received significantly more contact aggression than other age classes. Only the adult females increased aggression after the introduction of new animals. In study II, new social groups were formed and behavioral observations were made following formation. One group was formed from 7 pairs of familiar females and an unfamiliar adult male. This group had a high frequency of aggression during the first half hour, with contact aggression rising to peaks at 3 and 5 h. The second group consisted of 10 familiar females, 2 pairs of familiar females and an unfamiliar male. There was a significant peak in contact aggression 3 h into the observation.     

Factors affecting aggression among females in captive groups of rhesus macaques (Macaca mulatta)

Captive groups of primates often exhibit higher rates of aggression than wild, free-ranging groups. It is important to determine which factors influence aggression in captivity because aggression, particularly intense aggression, can be harmful to animal health and well-being. In this study, we investigated the effect of ground substrate as well as season, rank, age, and group size on rates of agonistic interactions per female in seven captive groups of rhesus macaques (n = 70 females, 1,723 focal samples) at the California National Primate Research Center. Agonistic interactions were divided into three categories: displacements, mild aggression, and intense aggression. Females living in enclosures with gravel substrate were 1.7 times more likely to be involved in intense aggression (e.g. chases and physical contact) than females living in enclosures with grass (Poisson regression model: P < 0.001). High-ranking females were at least 1.3 times more likely to be involved in mild (e.g. threats and lunges) aggression than lower-ranking females (low rank: P = 0.03; mid rank: P = 0.001). Females of all ranks were 1.5-1.9 times more likely to be involved in both intense and mild aggression during the breeding season than other seasons. Age and group size did not affect rates of mild or intense aggression. These findings indicate that although some aggression appears to be natural and unavoidable, i.e. aggression during the breeding season, the well-being of captive macaques can be improved by developing grass substrate in outdoor enclosures.     

Facility design for Bachelor Gorilla groups

This paper emerged from discussions following the Bachelor Gorilla 2000 Workshop Facility Design sessions. Although many ape facility design features are common to all gorillas, the purpose of this paper is to provide an overview of special needs of bachelor gorillas. For example, managing escalating aggression between maturing or silverback males may require a high degree of caregiver intervention and thus easy access to gorillas both on- and off-exhibit is beneficial. Facility design features that reduce contact aggression, increase affiliation, provide visual barriers or escape routes, and allow safe outlets for species-typical behavior are essential to the successful management of bachelor gorilla groups in zoos. Such provisions permit caregivers the ability to constantly monitor and modify the zoo environment to meet rapidly changing husbandry needs. Creatively designed habitats and continuing research on bachelor group dynamics will undoubtedly allow zoo managers to maintain all gorilla groups in an environmental and social context that enhances well-being over the long term. Zoo Biol 28:144–162, 2009. © 2009 Wiley-Liss, Inc.     

Behavioral changes across the menstrual cycle in isosexual groups of bonnet macaques (Macaca radiata)

Behavioral changes were systematically recorded across menstrual cycles over a six-month period in two laboratory-housed isosexual triads ofMacaca radiata (bonnet macaque). The purpose of this study was to determine whether females of this species demonstrate premenstrual behavioral changes as reported for humans, or heightened levels of aggression during the first half of their cycle as reported in some species of Old World monkeys. A total of 34 menstrual cycles were divided into five segments including two follicular, one periovulatory, and two luteal phases. The mean frequencies of behaviors were analyzed according to phase, rank, and time since triad formation, for a total of 34 menstrual cycles. There was no evidence of an increase in spatial separation during the premenstrual phase or during any other phase of the menstrual cycle. Although contact aggression did not show an increase during the early follicular half of the cycle (phases 1 and 2) or during the phase of the cycle immediately preceding menses (phase 5), contact aggression did show two peaks: one in the early-mid luteal half of the cycle (phase 4) and a peak in the mid-late follicular (phase 2). The non-aggressive hierarchical behaviors did not follow the same pattern as contact aggression. Instead the distribution of these behaviors showed a pattern similar to that of estrogen levels across the cycle. Subjects’ location in pen also varied significantly according to cycle phase: subjects spent more time on perches during the premenstrual phase and more time on the pen floor during the periovulatory phase. The increase in contact aggressive behaviors during the early-mid follicular phase and the mid-late luteal phase does not suggest a simple hormonal correlate as these two phases are characterized by high levels of estrogen and progesterone, respectively. However, the distribution of non-aggressive hierarchical behaviors suggest that this category of agonistic interaction may be related to mating competition among females of this species. Results are discussed with reference to the social behavior and promiscuous mating strategy ofM. radiata. The findings in this present study are compared with previous studies utilizing other species of Old World monkeys. Differences in study design and group composition are considered as factors effecting discrepant results both within and between species.     

Distraction sneakers decrease the expected level of aggression within groups: a game-theoretic model

Hawk-dove games have been extensively used to predict the conditions under which group-living animals should defend their resources against potential usurpers. Typically, game-theoretic models on aggression consider that resource defense may entail energetic and injury costs. However, intruders may also take advantage of owners who are busy fighting to sneak access to unguarded resources, imposing thereby an additional cost on the use of the escalated hawk strategy. In this article we modify the two-strategy hawk-dove game into a three-strategy hawk-dove-sneaker game that incorporates a distraction-sneaking tactic, allowing us to explore its consequences on the expected level of aggression within groups. Our model predicts a lower proportion of hawks and hence lower frequencies of aggressive interactions within groups than do previous two-strategy hawk-dove games. The extent to which distraction sneakers decrease the frequency of aggression within groups, however, depends on whether they search only for opportunities to join resources uncovered by other group members or for both unchallenged resources and opportunities to usurp.     

The fertility of daughters in common marmoset (Callithrix jacchus jacchus) family groups

Observations previously made in the field of inter-group mating raised questions concerning the fertility of daughters in family groups and about possible competition between mothers and daughters. To test whether daughters in captive family groups could become pregnant, daughters from four groups were placed with a strange male for a period of 30 min, twice a week for eight weeks. No changes were observed in the mother/daughter interactions during the time in which the daughters were allowed contact with the strange male. Two of the daughters showed signs of ovulation and the male only copulated with these animals. One daughter became pregnant, but it was aggression with her siblings rather than her mother which necessitated removing her from her family group.     

A comparison of aggressive play and aggression in free-living baboons, Papio anubis.

A quantitative comparison is given of aggressive play and aggression in terms of the duration and type of movement patterns employed, and the relationships between individuals. The mean length of contact in play was greater than in aggression, whereas chases tended to be longer in adult male aggression than in play, Linear hierarchies describing the overall direction of interactions between individuals were divisible into two sorts, according to the frequency with which roles were reversed. The regions of the body bitten and the proportions in which six types of movement pattern were used were related to the type of hierarchy involved. The results are discussed with reference to the possible functions of social play in the development of movement patterns and individual relationships.     

Postconflict Behavior of Spectacled Leaf Monkeys (Trachypithecus obscurus). II. Contact with Third Parties

We studied the postconflict interactions with group members other than the former opponent in two groups of spectacled leaf monkeys. We found no evidence of redirection of aggression towards other group members. Victims and aggressors sought affiliative contacts with uninvolved third parties. There was no evidence for consolation—affiliative contact initiated by an uninvolved third party, directed towards victims of aggression—in either group when all affiliative behaviors were considered. However, embracing was a characteristic first-contact interaction between individuals involved in aggression and third parties. This finding mirrors the results concerning reconciliation in spectacled leaf monkeys. Accordingly, embracing may be a true consolatory behavior in this species. When contacts with third parties occurred before or in the absence of reconciliation, the timing of such contacts fell within the time window during which reconciliation normally occurs. These contacts also resulted in affiliation levels twice that of baseline levels, supporting the idea that these contacts may function as a form of substitute reconciliation. We discuss these results in light of recent theories concerning postconflict behavior in primates.     

Semi-natural and enclosed groups of Macaca mulatta: a behavioral comparison

Differences in aggression and grooming frequencies between enclosed and semi-natural groups of Macaca mulatta were studied quantitatively. Group composition, group densities and time of day were held constant. Results indicated that semi-natural monkey groups were significantly more aggressive, while enclosed groups exhibited significantly higher grooming frequencies. These results raise the question: Do alterations in behavioral frequencies lead to changes in social behavior patterns and group social organization?     

Social interactions and aggression among male Madagascar hissing cockroaches (Gromphadorhina portentosa) in groups (Dictyoptera: Blaberidae)

We studied agonistic interactions among male Madagascar hissing cockroaches,Gromphadorhina portentosa, in groups of five (low-density) or 10 (high-density) males. Consistent with previous studies of male pairs, we observed aggression (Abdomen Flick, Abdomen Push, Butt, Lunge), submission (Crouch, Retreat), and noncontact behavior (Abdominal Extension, Abdomen Thrash, Agonistic Hiss, Stilt). Males at both densities performed all acts. However, males in the high-density group performed Abdomen Push significantly more often than males at a low density. The rate of each remaining act was unaffected by density. Regardless of density, males within social groups varied in aggression. More aggressive males utilized frontal assaults (Butt and Lunge) during interactions, while males displaying lower levels of aggression preferentially used the abdomen during interactions. More aggressive males performed Abdomen Flick more frequently, while males displaying lower levels of aggression performed Abdomen Push. We also investigated the relationship between male aggression and the four noncontact behaviors. We found that Abdominal Extension, Abdomen Thrash, and Agonistic Hiss were positively correlated with our aggregate measure of male aggression suggesting these are aggressive displays. Stilt was positively correlated with measures of both aggression and submission, leaving its function unclear. None of the behavioral acts examined in this study were highly correlated with male weight. Our results are discussed in light of possible hypotheses addressing the function of specific behavior during male-male competition.     

Coping with social tension: Sex differences in the effect of food provision to small rhesus monkey groups

In previous research, male chimpanzees (Pan troglodytes) were found to be more ‘conciliatory’ than females, in that after aggressive incidents males more often engaged in socially positive contact with former opponents. The first part of the present paper presents similar results for a large breeding troop of rhesus monkeys (Macaca mulatta). To investigate further the possible sex difference in postconflict behaviour, a series of experiments was carried out on small isosexual groups of young rhesus monkeys, three male and three female groups. Food competition was created by providing either a single apple piece (Monopoly test) or a handful of small pieces (Equality test). Observations lasted for 30 min, and behaviour in tests without food provisioning was also recorded. The predicted response to food was an increase in aggression followed by restorative behaviour, such as grooming. In Equality tests both sexes showed similar responses, i.e. increased aggression and decreased grooming and cohesiveness. In Monopoly tests, however, their responses differed. After the aggression increase, which occurred in all groups, males showed a significant increase in grooming and cohesiveness, whereas females showed the (non-significant) opposite trend. Another sex difference was that changes in grooming and aggression frequencies were related to the rank order in female groups, but not in male groups. Since there was no evidence for a direct causal connection between grooming and aggression, a new model is introduced which links grooming behaviour to the type of food provision rather than to the social disturbance by aggression. According to this model inequality in food distribution causes social tensions. Males actively try to reduce these tensions, whereas females do not. Some alternative explanations are also discussed.     

Female genital swellings have no effect on chimpanzee agonism in well-established groups

Reducing the incidence of aggression and wounding is a major concern for those managing socially housed chimpanzees (Pan troglodytes). Competition among adult males for access to sexually receptive females is believed to be one factor contributing to undesirably high levels of agonism. An observational analysis of two chimpanzee groups (n = 9, n = 8) was initiated to study this phenomenon. All-events sampling was used to record agonistic and sexual behavior on days when one female in the group showed a maximal genital swelling and on other days when no female in the group had a tumescent swelling. The result of the multivariate analysis of variance indicated that the presence of a female with a tumescent swelling had no effect on bluff displays or aggressive, submissive, or reconciliatory behavior but that sexual behavior was significantly increased. These results may be explained by the fact that long-term stability in group membership may reduce male–male competition for sexual access to females and the sometimes associated aggression and wounding. Colony managers dealing with well-established groups of chimpanzees should invest more of their effort in controlling aggression elicited by factors other than the cycling of a single female group member.     

THE ROLE OF MULTILEVEL SELECTION IN THE EVOLUTION OF SEXUAL CONFLICT IN THE WATER STRIDER AQUARIUS REMIGIS

In evolution, exploitative strategies often create a paradox in which the most successful individual strategy “within” the group is also the most detrimental strategy “for” the group, potentially resulting in extinction. With regard to sexual conflict, the overexploitation of females by harmful males can yield similar consequences. Despite these evolutionary implications, little research has addressed why sexual conflict does not ultimately drive populations to extinction. One possibility is that groups experiencing less sexual conflict are more productive than groups with greater conflict. However, most studies of sexual conflict are conducted in a single isolated group, disregarding the potential for selection among groups. We observed Aquarius remigis water striders in a naturalistic multigroup pool in which individuals could freely disperse among groups. The free movement of individuals generated variation in aggression and sex‐ratio among groups, thereby increasing the importance of between‐group selection compared to within‐group selection. Females dispersed away from local aggression, creating more favorable mating environments for less‐aggressive males. Furthermore, the use of contextual analysis revealed that individual male aggression positively predicted fitness whereas aggression at the group level negatively predicted fitness, empirically demonstrating the conflict between levels of selection acting on mating aggression.     

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.     

Dominance and aggression in social groups of male and female rats

Two experiments were performed to examine aggression and dominance in domestic male and female $\text{Rattus norvegicus}$ living in small mixed-sex (3 males and 3 females) groups. Experiment 1 examined the development of aggression in females. A single female (alpha) within each of the six colonies tested showed the preponderance of attacks on male intruders placed into the home-cage when male colony residents were absent. Over 12 weeks of intruder-aggression training female alphas showed only a mild nonsignificant elevation of aggressive behavior. A comparison of aggression of male and female colony alphas tested with opponents of each sex revealed that aggression was mainly directed at like-sex opponents, and that female attack was more defensive in character than male attack regardless of opponent sex. The highest intensity of aggression occurred when male alphas confronted male intruders. Although intruders never showed offense toward male residents, 61% of intruding males showed offense in response to attack by females.Experiment 2 investigated the relationship between aggressive dominance and competitive measures of dominance within each of 10 mixed-sex colonies. Alpha stat s of male and female colony residents did not reliably predict priority of access to food or water in tests of direct resource competition with like-sex colony members. When colony males were simultaneously tested for copulation, the copulatory behavior of alpha males was significantly greater than that of other colony males. Results are discussed in relation to the role of aggression in the reproductive strategy of male and female $\text{Rattus norvegicus}$.     

Behavioral and hormonal correlates of social dominance in stable and disrupted groups of male domestic fowl

An experiment was conducted to determine the behavioral and hormonal correlates of social dominance in stable and disrupted groups of male domestic fowl (Gallus domesticus). In addition, the relative importance of visual and auditory cues in the maintenance of dominance status in this species was assessed. Alpha roosters were removed from their six-bird home pens for 1 week, and placed in either (1) and adjacent pen in visual contact with penmates, (2) and adjacent pen where visual, but not auditory, contact was prevented, or (3) a different building where they were in visual and auditory isolation. Dominance rank, aggressive activity, and crowing frequency were intercorrelated during the premove period. There was a significant increase in both aggression and crowing among the males remaining in the home pen following removal of the alpha male, except in the treatment in which males were still in visual contact with the removed alpha, in which only aggression increased. Increases in crowing frequency were also noted in alpha males. High levels of aggression occurred when alpha males were returned except in those pens in which visual contact had been maintained. Visual cues thus appear to be more important than auditory cues alone with respect to the maintenance of dominant social status in roosters. There were no clear correlations between behavioral variables and plasma androgen or corticosterone levels in either the stable or disrupted flocks, and no consistent changes in the levels of these hormones as a result of experimental manipulation; several possible explanations for this finding are discussed.     

Weaponry,size, and sex ratio affect spatial distribution within small and large groups of the maritime earwig (Anisolabis maritima)

Dispersion patterns within a group can reveal important aspects about social interactions and sexual selection within a species. We examined the distribution patterns of the maritime earwig (Anisolabis maritima), an insect well suited for studies of aggression, sociality, and sexual selection since both sexes live in close proximity and possess weaponry in the form of sexually dimorphic pincers. To examine intra‐ and intersexual interactions within small groups, we conducted trials with three earwigs with limited access to shelters. In single‐sex trios, we found that both males and females exhibited strong size‐based intrasexual aggression, as larger individuals were less likely to be excluded from shelters; however, males were more likely to cohabitate than females. In mixed‐sex trios, we found that both males and females preferred smaller opposite‐sex partners, and cohabitation patterns indicate that both sex‐ and size‐based differences in aggression can influence overall spatial distribution. We also examined larger single‐sex and mixed‐sex groups of 18 earwigs to determine whether they had random, uniform, or clumped distributions. Similar to previous field observations, males tended to form aggregations, whereas females were distributed uniformly, a pattern indicative of territoriality. Mixed‐sex groups, on the other hand, were uniform during nocturnal periods of high activity but then become clumped after settling into more stable daytime positions. Overall, our results suggest that females have high levels of aggression regardless of the social context, whereas males alter their aggressive behavior in the presence of females.     

Social rank and habitat use in willow tit groups

One view depiets dominance and subordination as roles with equal fitness played as parts of a mixed behavioural strategy, but subordination can also be maintained as a transient strategy of ‘hopeful dominants’. Resource access in groups of non-kin willow tits, Parus montanus, was related to social rank. Dominants excluded subordinates from the preferred microhabitat. Fitness in terms of survival was correlated with access to the preferred foraging sites, indicating that a mixed-strategy explanation is inadequate. Social rank was linked to age, and first-year subordinates behaved as predicted for hopeful dominants. They were not ‘herded’ by dominant group mates because subordinates actively maintained contact with dominants. One possible reason for subordinates joining winter groups might be the opportunity to reduce dominants' pay-off from aggression.     

Aggression in pigtailed macaque (Macaca nemestrina) breeding groups affects pregnancy outcome

Past research has shown that aggressive behaviors can affect female reproductive outcome in nonhuman primate captive breeding programs. In this study, aggressive behaviors were recorded in a colony of pigtailed macaque monkeys (Macaca nemestrina) and related to pregnancy outcome. For 22 weeks, behavioral data were collected from nine breeding groups, consisting of zero to one male (some males were removed after a cycle of conceptions for husbandry reasons) and four to eight females. Observations included all occurrences of 11 aggressive behaviors during 15 min observation sessions, 1-3 times a week. Mean weekly aggression levels during the study period were determined for each group as well as for each pregnancy. Aggression data were summarized with Principal Components Analyses. Results indicate that pigtailed macaque aggression falls into five distinctive categories: warn, engage, threaten, pursue, and attack. Breeding groups differed in their levels of aggression, even after controlling for group size, presence of a sire, and group stability. Levels of the five aggression categories were found to affect the probability that a pregnancy ended in either a natural birth of a live infant, a clinical intervention producing a live infant, or a nonviable outcome. The predictive value of aggression was significant when clinical interventions were included as possible reproductive outcomes. Behavioral observation of captive groups could identify "risk" conditions affecting pregnancy outcome and the requirement for clinical intervention.