Effects of group composition on agonistic behavior of captive pigtail macaques,Macaca nemestrina

We surveyed agonistic behaviors of 20 captive groups of pigtail macaques (Macaca nemestrina) housed under identical spatial conditions. Fifteen groups contained one male each; the other five groups contained no adult males. Groups included six to twelve adult females, some of which had infants with them. We found no relationship between social density of groups and incidence of agonistic behavior, but significantly more contact aggression (grab, hit, push, bite) and noncontact aggression (chase, open-mouth “threat,” bark vocalization) occurred among females in groups containing no males than in those containing one male each. Apparently, males played an important role in the inhibition of intragroup conflict. We also found that females in groups containing males exhibited less noncontact aggression if infants were present than if no infants resided in their groups. Thus, competition of females over infants must not have been an important constituent of intragroup conflict under the conditions of this survey.     

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.     

Intergroup Relationships in Western Black-and-White Colobus, <Emphasis Type="Italic">Colobus polykomos polykomos</Emphasis>

We investigated patterns of intergroup relationships in western black-and-white colobus, Colobus polykomos, in Taï National Park, Côte d'Ivoire, between 1993 and 1999. They live in one-male multifemale units, and demonstrate male dispersal and occasional dispersal by females. Solitary males and all-male bands are absent or very rare. Our aim was to investigate the function of female and male aggression during intergroup interactions. The species is particularly interesting because, in contrast to predictions from socioecological models, female aggression occurs during intergroup interactions in combination with female dispersal. Home ranges of neighboring groups overlapped considerably and groups lacked an area of exclusive access. Intergroup interactions occurred once every 6.6 observation days. Encounters were either peaceful (12%), or involved displays and threats (25%) or chases and fights (63%). Females interacted in 74% and males in 98% of aggressive intergroup encounters. We found little to no indication that male and female aggression correlated with the presence of food, importance of a location, or presence of infants or receptive females. However, females were more often aggressive during the months when the group depended strongly on seeds from Pentaclethra macrophylla. We also observed forays by males to other groups. Forays occurred on average once every 20 observation days. In 75% of the forays, the intruding male chased members of the target group. In 25% of the forays 1–3 females joined their male but females never attacked the target group. Our main study group was the target of such forays significantly more often when young infants were present in the group than when not. We conclude that female aggression between groups was related to food procurement and that male forays might be related to infanticide.     

Fighting and Mating Between Groups in a Cooperatively Breeding Mammal, the Banded Mongoose

Many cooperatively breeding animals actively defend a territory containing resources such as food and shelter, which are essential for reproduction. Some observations, however, indicate that conflicts between groups are often triggered by the attempts of males or females, or both, to gain extra‐group copulations. We studied interactions between 12 groups of banded mongooses (Mungos mungo) in Uganda to test whether the frequency of inter‐group encounters was linked to the reproductive status of females, and conducted an experiment to examine the responses of individuals to mongooses from other groups. The rate at which focal groups fought with other groups was higher when its females were in estrous, suggesting that many fights take place over access to mates. Both males and estrous females were instrumental in instigating encounters with rival groups, and extra‐group copulation in the midst of a fight was observed on three occasions. We experimentally simulated encounters with foreign individuals by presenting each of six groups with cages containing a male and a female from a rival group. Subordinate males were the first to investigate these ‘intruders’, spent more time around the intruders' cages, and were more aggressive to the intruders than either dominant males or females. Subordinate males directed more attention and aggression towards the male intruder than the female intruder. We suggest that male banded mongooses actively seek extra‐group copulations in pursuit of paternity, while females may actively seek extra‐group copulations as a way of reducing inbreeding depression.     

Sex differences and developmental effects of oxytocin on aggression and social behavior in prairie voles (Microtus ochrogaster)

Various hormones, including sex steroids and neuropeptides, have been implicated in aggression. In this study we examined (1) sex differences in intrasexual aggression in na?ve prairie voles; (2) the effects of developmental manipulations of oxytocin on intrasexual aggression; and (3) changes in patterns of intrasexual aggression after brief exposure to an animal of the opposite sex. Within 24 h of birth, infants were randomly assigned to receive either an injection of oxytocin (OT) or oxytocin antagonist (OTA) or to one of two control (CTL) groups receiving either isotonic saline or handling without injection. As adults, animals were tested twice in a neutral arena; before (Test 1) and 24 h after (Test 2) a 4-h exposure to an animal of the opposite sex. In Test 1, CTL males were more likely to show aggressive and less likely to show social behavior than CTL females. No significant treatment differences were observed within either sex in Test 1. In Test 2, after brief exposure to a male, females treated with OT became more aggressive and less social than OTA or CTL females. Male aggressive behavior did not change after exposure to a female. An increase in aggression and decline in social behavior toward other females, seen here in OT-treated females, is typically observed only following several days of female-male cohabitation. These findings demonstrate a sex difference in intrasexual aggression and suggest that neonatal exposure to OT may facilitate the onset of the mate-guarding component of pair bonding in female prairie voles.     

Female competition in chimpanzees

Female chimpanzees exhibit exceptionally slow rates of reproduction and raise their offspring without direct paternal care. Therefore, their reproductive success depends critically on long-term access to high-quality food resources over a long lifespan. Chimpanzee communities contain multiple adult males, multiple adult females and their offspring. Because males are philopatric and jointly defend the community range while most females transfer to new communities before breeding, adult females are typically surrounded by unrelated competitors. Communities are fission–fusion societies in which individuals spend time alone or in fluid subgroups, whose size depends mostly on the abundance and distribution of food. To varying extents in different populations, females avoid direct competition by foraging alone or in small groups in distinct, but overlapping core areas within the community range to which they show high fidelity. Although rates of aggression are low, females compete for space and access to food. High rank correlates with high reproductive success, and high-ranking females win direct contests for food and gain preferential access to resource-rich sites. Females are aggressive to immigrant females and even kill the newborn infants of community members. The intensity of such aggression correlates with population density. These patterns are compared to those in other species, including humans.     

The Payoff of Fighting in House Crickets: Motivational Asymmetry Increases Male Aggression and Mating Success

Motivational asymmetry caused by differences in subjective resource value is a key component of strategic models of aggression. We study the role of motivational asymmetry in determining differential aggression and mating success of male house crickets, Acheta domesticus. We also assess the extent to which mating differences associated with motivational asymmetry are due to direct male–male fighting vs. male–female interactions. We manipulated male motivation to compete for a mating opportunity by providing males with either no access or nightly access to females for 4 d prior to the experiment. As predicted, when males from each treatment had to compete for the female, those with lower prior access were more aggressive and mated more often. In contrast, when males from each treatment were paired individually with females, there was no significant difference in the frequencies with which they pursued, courted or mated with females. We also found no evidence for female choice based on motivational asymmetry; the rate of successful courtship did not differ between treatments. We conclude that prior mate encounter rate can generate motivational asymmetry, leading to differential mating success mediated by direct male–male aggression.     

Bystanders affect the outcome of mother–infant interactions in rhesus macaques

Animal communication involves the transfer of information between a sender and one or more receivers. However, such interactions do not happen in a social vacuum; third parties are typically present, who can potentially eavesdrop upon or intervene in the interaction. The importance of such bystanders in shaping the outcome of communicative interactions has been widely studied in humans, but has only recently received attention in other animal species. Here, we studied bouts of infant crying among rhesus macaques (Macaca mulatta) in order to investigate how the presence of bystanders may affect the outcome of this signalling interaction between infants and mothers. It was hypothesized that, as crying is acoustically aversive, bystanders may be aggressive to the mother or the infant in order to bring the crying bout to a close. Consequently, it was predicted that mothers should acquiesce more often to crying if in the presence of potentially aggressive animals. In line with this prediction, it was found that mothers gave infants access to the nipple significantly more often when crying occurred in the presence of animals that posed a high risk of aggression towards them. Both mothers and infants tended to receive more aggression from bystanders during crying bouts than outside of this time, although such aggression was extremely rare and was received by less than half of the mothers and infants in the study. Mothers were also found to be significantly more aggressive to their infants while the latter were crying than outside of crying bouts. These results provide new insight into the complex dynamics of mother–offspring conflict, and indicate that bystanders may play an important role in shaping the outcome of signalling interactions between infants and their mothers.     

Family Feuds: Severe Aggression among Cooperatively Breeding Cotton-Top Tamarins

A survey of severe aggression occurring over the 20 year history of a cotton-top tamarin (Saguinus oedipus) colony indicated that most aggression was between same-sex individuals, with fights among siblings more common than parent-offspring conflict. Males engaged in severe aggression more often than females did, paralleling the dimorphic aggressive response to same-sex intruders in this species. Female-female aggression occurred at larger group sizes than male-male aggression, suggesting a higher threshold for female aggression. Much sibling aggression was directed by postpubertal subadults toward prepubertal siblings. Only 17% of aggression occurred in association with infant births, and these cases rarely involved parents and offspring suggesting that aggression was not used by reproductive animals to defend their breeding position. Severe aggression was often associated with attempts to introduce stepparents or movement of individuals in and out of the group. The mean group sizes when aggression is observed in captivity are close to the maximum group sizes observed in wild tamarins, which suggests that severe aggression in captive groups may reflect processes related to dispersal in wild tamarins.     

Population density and the evolution of male aggression

In some cases male animals engage in aggressive contests for access to females, in others they adopt more passive strategies and invest in traits that assist them in detecting females or in competing with rivals in other ways, such as sperm competition. One possible factor determining the fitness of these different strategies is population density. Theoretically, aggressive tactics should be found at intermediate population densities. At low densities males that invest in traits related to searching for mates could be favoured, whereas at the highest densities males that fight over females might pay excessive costs for this behaviour because of the number of rival males that they will encounter. Current empirical evidence is mostly consistent with this scheme: in some cases it seems that traits that are associated with locating mates are favoured at low densities, with aggression related traits favoured at higher densities, and in other cases aggression is selected but as density increases less aggressive strategies become more common. There remain substantial differences between species, however, and I discuss how variation in mating system, in the costs of aggression and in the nature of sperm competition, plus ecological differences between species, can change the relationship between population density and the fitness consequences of aggressive and passive behavioural strategies.     

Condition-dependent reproductive tactics by large and small anadromous male sockeye salmon Oncorhynchus nerka

Reproductive tactics, aggression and courtship by large and small (jack) male sockeye salmon Oncorhynchus nerka with ripe females in temporary enclosures depended on the size and behaviour of the competitor, not the size of the focal male. When paired with other jacks or with large but less aggressive males, jacks fought for access to ripe females rather than sneaking, but used sneaking tactics when paired with large, aggressive males. Dominant males of both size groups courted actively and females spawned readily with all combinations of males.     

Sex differences in the behaviour of immature captive lowland gorillas

Studies of the behaviour of 26 (12 males and 14 females) captive infant and juvenile lowland gorillas showed clear sex differences. Females showed greater interest in young infants and were more active in nest building as well as in solitary and social grooming. Males were more active in locomotive, dominance, and aggressive behaviour and in social play. Hand-rearing further increased aggression. Males were more aggressive when they lived with only one partner, and they rose in rank even above older females, a pattern that has not been observed in naturally reared gorillas.     

Aggression and worker control of caste fate in a multiple-queen wasp, Parachartergus colobopterus

Although famously cooperative, social insect colonies harbour considerable potential for genetic conflict among colonymates. This conflict may be expressed behaviourally as aggression by workers. We investigated aggression in 34 colonies of the wasp Parachartergus colobopterus, by evaluating the characteristics of both instigators and victims of aggressive interactions. We estimated genetic relatedness and queen number using DNA microsatellites and found that workers and emerging females should be most in conflict over the caste of the latter when there are many queens on the nest. We found that aggressive interactions are more likely to involve older workers attacking either males or younger workers, and that victim and aggressor females have more ovarian development than randomly sampled colonymates. Moreover, mated females with low levels of ovarian development relative to active queens were also more likely to be aggressors and victims than were randomly sampled females. Aggression among females supports the hypothesis that older workers use aggression towards younger females as a means of policing the development of emerging females into queens. Workers also may use aggression to suppress the reproduction of some mated females. Our findings thus support the hypothesis that genetic conflicts of interest motivate worker aggression in swarm-founding wasp colonies.     

Sexual mimicry in hyenas

Genital masculinization in female spotted hyenas has been widely explained as an incidental consequence of high androgen levels. High androgen levels, in turn, were supposed to be favored because they led to adaptive aggressive behavior. Incidental androgenization is no longer a tenable hypothesis, however, because genital masculinization has been shown to proceed in the absence of androgenic steroids. Thus, an alternative hypothesis is required. The genitals of spotted hyena females are not simply masculinized, but exhibit a detailed physical resemblance to the male genitalia. In the absence of satisfactory alternative explanations, we propose that selection may have favored sexual mimicry in females because they are more likely than males to be targets of aggression from other females. Male-like camouflage could theoretically be protective in three contexts: neonate sibling aggression, infanticide by conspecific females, and interclan territoriality. Current data suggest that if sexual mimicry is important, its effects are strongest among infants.     

Natural genetic variation in social niche construction: social effects of aggression drive disruptive sexual selection in Drosophila melanogaster

Social niche construction (SNC) occurs when animals actively shape their social environments. Currently the fitness consequences of SNC are poorly understood, and no study has examined whether variation in SNC has a genetic basis. Here we report the first instance of genetic variation in SNC by showing that Drosophila male aggression shapes the social environment. We allowed flies of different genotypes to interact in complex arenas; we measured the number and sex of individuals in the groups that formed and counted instances of mating. Arenas containing more aggressive male genotypes formed groups with fewer males, demonstrating that aggressive male genotypes experienced different social environments than nonaggressive genotypes. Further, genotypes with highest mating success were those whose SNC behavior generated the social environment in which they were most adept at mating: genotypes who mate most often after winning aggressive encounters benefit from aggressive SNC, while genotypes who mate most often after losing achieve high mating rates by forgoing aggression. The presence of these alternative strategies-which were robust across eight population densities-revealed that selection on aggression and context-dependent mating was disruptive, consistent with the hypothesis that SNC can maintain genetic variation in multiple behaviors.     

Successive aggression: another pattern of polyadic aggressive interactions in a captive group of Japanese macaques

Several patterns of polyadic aggressive interactions have been previously reported. Here, we describe another pattern of polyadic interactions in a captive group of Japanese macaques (Macaca fuscata fuscata) that we designate as "successive aggression". We defined successive aggression as aggression by the original actor toward a second recipient within a very short time interval after the initial aggression toward the first recipient. We compared the patterns and characteristics of successive aggression to those of redirection. Among 2,698 recorded aggressive interactions, 80 involved successive aggressions and 75 were classified as redirections. Females, especially adult females, performed and received more successive aggression, whereas males, especially adult males, performed and received more redirection. Successive aggression often occurred when the first recipient exhibited counter-aggression. Successive aggression was then directed toward an individual related to the first recipient, such as the mother, offspring or sibling. The targets of redirection were not relatives of the first recipient in most cases, but were clearly subordinate individuals. The dominance relationships among the aggressor, the first recipient and the second recipients were usually non-linear for successive aggression, but were linear for most cases of redirection. These results suggest that monkeys can anticipate possible opponents who may intervene in ongoing aggressive interactions and suppress them, even though they are not yet hostile toward these individuals. Successive aggression may function to establish and maintain dominance relationships among matrilineal groups through repeated confirmations.     

Integration of New Males into Four Social Groups of Tufted Capuchins (Cebus apella)

We examined how aggressive, affiliative, and sexual behavior function to integrate male capuchins (Cebus apella) into a new social group. Nine males were exchanged among four social groups. We performed instantaneous scans and all-occurrence sampling during baseline, introduction, and follow-up periods. The study included three different introduction situations: 1) males familiar to one another were introduced to a group with no other adult male, 2) males unfamiliar to one another were introduced to a group with no other adult male, and 3) males familiar to one another were introduced to a group with an existing elderly, resident male. Severe aggression occurred in situations 2 and 3, but the introductions were peaceful in situation 1. In all cases proceptive females were among the first individuals to affiliate with the males, and males did not appear to compete for access to proceptive females. Following their period of proceptivity, the females that had cycled remained preferred social partners for the males. Immature animals also quickly affiliated with the new males, and the males tolerated the attention from immatures. Affiliative relationships between the males and nonproceptive females developed slowly, and while male-female aggression was mild, aggression among adult males (familiar and unfamiliar) had the potential to be severe.     

Sex and Weapons: Contrasting Sexual Dimorphisms in Weaponry and Aggression in Snapping Shrimp

Sexual dimorphisms in weaponry and aggression are common in species in which one sex (usually males) competes for access to mates or resources necessary for reproduction – sexually dimorphic weaponry and aggression, in other words, are frequently the result of intrasexual selection. In snapping shrimp, the major chela (snapping claw) can be a deadly weapon, and males of many species have larger chelae than females, a pattern readily interpreted as resulting from intrasexual selection. Thus, males might be expected to show more sex‐specific aggression than females, and be more aggressive overall. We tested these predictions in two species of snapping shrimp in a territorial defense context. Neither of these predictions was supported: in both species, females, but not males, engaged in sex‐specific aggression and females were more aggressive than males overall. These contrasting sexual dimorphisms – larger weaponry in males but higher aggression in females – highlight the importance of considering the function of weaponry and aggression in contexts other than direct competitions over mates. In addition, species differences in the degree of sexual dimorphism in chela size were due to differences in female, not male, chela size, and the species with greater sexual dimorphism in weaponry was significantly less aggressive overall; also, while paired and solitary males did not differ in residual chela size, for the species with greater sexual dimorphism, females carrying embryos had smaller residual chela sizes. These results suggest that understanding the sexual dimorphisms in weaponry and aggression in snapping shrimp requires understanding the relative costs and benefits of both in females as well as males.     

Post-conception reproductive competition in cooperatively breeding common marmosets

Common marmosets are cooperatively breeding monkeys that exhibit high female reproductive skew. Subordinate females usually fail to breed as a consequence of ovulation suppression and inhibition of sexual behavior, and, even when they do breed, typically rear fewer infants than dominants. We evaluated possible mechanisms of post-conception reproductive competition by comparing hormonal profiles across pregnancy, pregnancy outcomes, infant survivorship, and behavior in laboratory-housed families containing one (N=9) or two (N=7) breeding females. Breeding females in plurally breeding groups did not exhibit well-defined dominance relationships and rarely engaged in escalated aggression with one another. No significant differences were found among singly breeding mothers, plurally breeding mothers, and plurally breeding daughters in urinary chorionic gonadotropin or estradiol sulfate concentrations during pregnancy, fetal biparietal diameter, frequency of spontaneous abortion, frequency of stillbirths, number of live-born infants per litter, or infant mortality rates. When females gave birth while another female in the family was pregnant, however, their infants were highly likely to be killed. The perpetrator was definitively identified in only one family, in which a pregnant female killed her daughter's infant. These results are consistent with observations of free-living common marmosets and suggest that breeding females do not regularly influence one another's pregnancy outcomes, but that they may commonly kill each other's infants, especially during their own pregnancy. Our findings further suggest that infanticide by breeding females may have selected for the evolution of reproductive restraint in subordinate female marmosets.     

Sex differences in intrasexual aggression among sex-role-reversed,cooperatively breeding cichlid fish <Emphasis Type="Italic">Julidochromis regani</Emphasis>

In sex-role-reversed species, females compete for resources (e.g., mates) more intensively than do males. However, it remains unclear whether these species exhibit sex differences in the intensity of aggressive behavior in the context of within-sex contests. Cichlid fish in the genus Julidochromis exhibit intraspecific variation in mating systems, ranging from monogamy to cooperative polyandry with sex-role reversal. In the study reported here, we observed aggressive interactions among three same-sex individuals in Julidochromis regani in the laboratory and tested whether inter-female aggression was more intense than inter-male aggression. Although difference in body size strongly determined the direction of aggression in fish, aggression by a smaller-sized individuals toward larger ones was occasionally observed. This type of aggression was common between individuals of a similar body size (≤5 mm) and occurred more frequently among females than males. In contrast, differences in body size and sex did not affect the frequency of aggression by larger-sized individuals against smaller ones. Bidirectional aggression (i.e., mouth fighting) occurred frequently when two individuals had similar body size, and there was no difference in its frequency between sexes. However, temporal analysis showed that females performed bidirectional aggression more persistently than males. These sex differences in the intensity of intrasexual aggression could be the behavioral mechanisms underpinning cooperative polyandry.