Reactions of nuclear-family–reared rhesus macaques to the births of younger siblings

A group of juvenile rhesus macaques (Macaca mulatta) living in a nuclear-family laboratory environment was studied to determine their responses to the births of siblings. The frequencies of interactions with family members (mothers, fathers, and new siblings) and nonfamily (peers, unrelated infants, and unrelated adults) were studied over both the year preceding and the year following sibling birth. The frequencies of specific behaviors in each of those interactions and the frequencies of interactions in each area of the nuclear-family unit (home cage, play area, or other families' cage) were also examined. After new siblings arrived, several measures of interactions with mothers, fathers, and new siblings increased significantly; by contrast interactions with peers decreased substantially over the post-birth year. Although the frequency of interactions in home cages remained stable over the 2-year period, interactions outside of the subjects' home cages decreased significantly after siblings were born. An additional subject group whose mothers became pregnant but failed to deliver viable offspring showed no significant changes in total levels of interactions with peers; they did, however, exhibit increases in some interactions with unrelated infants and adults. Female juveniles interacted with new siblings significantly more often than did males when siblings were less than 6 months old, but as siblings grew older (6–12 months), females' levels of interaction with them fell to a level equal to that of males. In the nuclear-family social structure, the birth of a sibling resulted in an increased emphasis on family interactions at the expense of peer interactions.     

Birth-season interactions of adult female Japanese Macaques(Macaca fuscata) without newborn infants

The affiliative interactions of 11 adult female Japanese macaques that did not deliver an infant during the 1981 birth season of the Arashiyama West troop were examined. Consideration was given to the effects of kinship as a structuring element in these birth-season interactions and to the degree of association with various categories of troop members based on age, sex, and (in the case of adult females) whether or not the females were new mothers. Females without infants interacted predominantly with their yearling off-spring, although it was the behavior of the offspring that precipitated the interaction. These females were active in soliciting affiliation with nonkin new mothers, whereas female matrilineal relatives with new infants approached and remained in proximity to them more than did nonrelated new mothers. Females without newborns groomed and approached nonkin infants more than infants within their own matriline, and these infants were predominantly those of females in the highest-ranking matriline of the troop. Adult males were responsible for 40% of all grooming received from nonkin by the females without newborns, and these males approached them significantly more than did other adult females without infants. These patterns demonstrate that the structure of social relationships is influenced by the particular dynamics of troop contexts such as birth seasons, as well as by enduring, broad-based affinities which are less affected by cyclic changes in troop context.     

Social behavior of infant and mother Japanese Macaques (Macaca fuscata): Effects of kinship,partner sex,and infant sex

The behavioral interactions of 22 infant and mother Japanese macaques with other group members were studied. Focal-animal observations were made from the time of each infant’s birth until 1 year of age. Infants and mothers both displayed exceedingly strong preferences for associating with matrilineal kin and, specifically, for female kin. The degree of genetic relatedness was positively correlated with levels of spatial proximity, contact, grooming, aggression, and play. Overall frequencies of interactions with nonkin were very low, and partner sex was not an important factor in interactions with nonkin. There were no significant differences between male and female infants in interactions with kin versus nonkin. There was only one significant difference between male and female infants in interactions with males versus females: female infants showed stronger preferences for initiating proximity with females over males than did male infants. Because mothers provide the focal point for infant interactions during the first year of life, we compared the behavior of infants and mothers. Mothers were the recipients of more social interactions than were infants, mothers engaged in more grooming than did infants, and infants engaged in more social play than did mothers. These findings are only partially consistent with kin-selection theory, and the inadequacies of studying matrilineal kin discrimination to test kin selection are reviewed. The near-absence of infant sex differences in associations with social partners suggests that although maternal kin other than the mother are important to infant socialization, they probably do not contribute to the development of behavioral sex differences until after the first year of life.     

Behaviours promoting grouping or dispersal of mother and neonates in ovoviviparous cockroaches

Summary. The nature of early relationships between mother and nymphs and among siblings was compared in four cockroach species belonging to the same ovoviviparous family (Zetoborinae) in order to characterise the behavioural interactions favouring dispersal or maintenance of the group of neonates. Behavioural interactions between mothers and their new-born nymphs and between two sibling neonates were video recorded and analysed with flow charts on factorial maps. In the solitary species Thanatophyllum akinetum, nymphs dispersed a few hours after birth without aggressiveness between siblings or between mother and offspring. In contrast females of Schultesia lampyridiformis displayed a behaviour never previously observed in cockroaches: aggression towards their own neonate nymphs, which could contribute to their dispersal. In Phortioeca nimbata and Lanxoblatta emarginata, nymphs stayed with their mother for 10 days, but social interactions differed between the two sp ecies: P. nimbata mothers actively searched for their nymphs, this behaviour being favoured by an active search for mutual contact by the nymphs themselves, while L. emarginata nymphs sought more actively the proximity of their mother but less contact between themselves. Thus, different species of Zetoborinae presented two types of dispersal of the young and two types of maintenance of the birth group, both achieved by specific behavioural interactions. The relevance of behavioural interactions for the characterisation of early gregarism and parental care is discussed.     

Socialization in spectacled parrotlets (Forpus conspicillatus): how juveniles compensate for the lack of siblings

  Socialization, that is, the process of the integration of an individual into the social system and the acquisition of basic social traits, has been studied in a variety of vertebrate species. Deprivation experiments have shown that the lack of early affiliative relationships has negative long-term effects on social behaviour and reproduction. Most studies emphasized the importance of the mother-infant relationship. However, it soon became evident that peer relationships to like-aged conspecifics, such as siblings and unrelated juveniles of nearly the same age, are also important for socialization. Whether individuals, deprived of one of these important social components, are able to compensate for this lack is not known. Here I tested the hypothesis that in spectacled parrotlets (Forpus conspicillatus), singletons (i.e., those that grew up without siblings) were able to compensate for this social lack by establishing and strengthening other relationships. In spectacled parrotlets socialization depends on the formation of long-lasting sibling groups. However, singletons without nest mates are quite common (about 6%), both in captivity and in nature. In this study I analysed the affiliative interactions with other group members of five singletons in comparison to eight siblings, from fledging to the 36th week after fledging, within a group of adult and juvenile conspecifics. In contrast to siblings, the singletons prolonged the relationships with their parents. Then they established a ”host sibling group” with other like-aged individuals. All singletons became well integrated into the group and established exclusive pair bonds. It is concluded that juvenile singletons of spectacled parrotlets are able to compensate for the lack of siblings by the development of alternative socialization tactics. Received: 12 December 1998 / Received in revised form: 17 May 1999 / Accepted: 7 June 1999     

Male-male relationships among wild bonobos (Pan paniscus) at Wamba,Republic of Zaire

Male-male relationships among wild bonobos (Pan paniscus) in two adjacent unitgroups (E1 and E2 groups), which were formed by division of the E group, were studied at Wamba, in the Central Zaire Basin, by analyzing the proximity and social interactions among males. Dominant-subordinate relationships between a male-male dyad were easily recognized from the directions of individual agonistic interactions. Male bonobos rarely joined forces in aggression. Clear differences in social status existed between adult and adolescent male bonobos in both groups, as reported in the case of chimpanzees (Pan troglodytes). The presence of mothers in the unit-group greatly influenced the dominant-subordinate relationships among males through strong mother-son bonds in both groups. However, the extent of the mother-son bonds differed between the groups. Males in the E2 group participated more frequently in agonistic or affinitive interactions than did males in the E1 group. Males in the E1 group were divided spatially into several clusters, while there were cohesive relationships among the adult males in the E2 group. The difference in intensities of mother-son bonds between the groups may be explained by the distribution of males at the time of the division of the E group. Differences in male-male relationships between bonobos and chimpanzees seem to be related to differences in intra- and inter-unit-group competition among males between the two species. Male chimpanzees may achieve coexistence by manipulating ambivalent relationships that are caused by intra- and inter-unit-group competition among them, while male bonobos may achieve coexistence by decreasing intra- and inter-unit-group competition among them.     

Sex differences in infant integration in a semifree-ranging group of Barbary macaques (Macaca sylvanus,L. 1758) at Salem,Germany

Sixteen multiparous Barbary macaque females with newborns were studied over a 16 month period within the context of their naturally formed group. Analysis of their social behavior revealed 1) triadic interactions involving focal females, their newborns, and other group members occurred mainly with other females; and 2) mothers with female newborns interacted mainly with females of their own matriline, while mothers with male newborns interacted mainly with nonmatriline females. Observed in two successive birth seasons, this pattern indicates that partners of maternal interactions chose each other according to the sex of the newborn. Measures of distance from the mother also reflected differences between infants of different sex. At about five months of age, female infants were observed close to their mothers significantly more often than males. This finding follows the pattern of a sex-specific infant socialization process which integrates female infants into the network of their matrilines and male infants into the broader group. This sex-specific integration pattern is interpreted as supporting female philopatry and male dispersal. © 1995 Wiley-Liss, Inc.     

The effects of litter-size variation on the development of play behaviour in the domestic cat: litters of one and two

In many litter-bearing species, litter-mates interact with each other in a variety of ways during early life. In the domestic cat, Felis catus, social play is a prominent form of interaction between litter-mates. The present study investigated how the lack of a litter-mate affected the development of kitten play and of the mother-kitten relationship. The subjects were 14 litters of domestic kittens living with their mothers in large indoor cages. Seven litters contained two male kittens, and the other seven contained single male kittens. Social interactions within the families were observed from day 22 to day 83 after birth. Single kittens played a more active part in maintaining close proximity to their mothers and directed more playful behaviour at them than did kittens with a sibling. Although single-kitten mothers avoided their offspring more than did mothers of litters of two, they also directed much higher levels of play behaviour at them. Despite the marked difference in the mother-kitten play relationship in the two litter types, single kittens experienced quantitatively less social play than did kittens with siblings. As the kittens grew older, single-kitten mothers showed higher levels of aggression towards their young than did mothers of litters of two.     

Intergroup affiliative interactions and intergroup transfer of young male Japanese macaques (Macaca fuscata)

The social interactions between young male Japanese macaques (Macaca fuscata) and members of two adjacent groups were studied. Young males usually associated with members of a single group. Although some young males occasionally interacted with members of the neighboring group, the frequency of their intergroup affiliative interactions was much lower than that of their intragroup interactions. The intergroup affiliative interactions were less symmetrical than the intragroup interactions. Three- or four-year-old males who remained in their natal group interacted with males of the neighboring group, whereas males over 5 years old did not. Young males revealed a dramatic change in their association partners from males in one group to those in another during the course of their intergroup transfer. Males who remained in their natal group did not attempt to interact with females of the neighboring group. In contrast, males who had transferred to a non-natal group interacted with females in their natal group. It is suggested that intergroup affiliative interactions and intergroup transfer of young male macaques are influenced by close associations between males. The immediate motivation for transfer of young natal macaques may be some attraction to males outside their group rather than sexual attraction to unfamiliar females.     

Intragroup Social Relationships of Male Alouatta palliata on Barro Colorado Island,Republic of Panama

We examined social and spatial relations of adult males in one group of mantled howling monkeys (Alouatta palliata) on Barro Colorado Island (BCI) in central Panama to document patterns of association. Beyond the existence of an alpha male, we could not distinguish any linear dominance hierarchy among the 6 study males. All males copulated with estrous females. Our findings contrast with reports of intragroup male behavior in Costa Rican howlers. Study males engaged in little or no affiliative or agonistic behavior with one another, but engaged in significantly more such interactions with females. The alpha male, the oldest male and a younger male were most frequently in association with females. Of group males, the oldest male associated significantly more with other males. Overall, male behavior in mantled howling monkeys on BCI generally followed the van Hooff and van Schaik (1994) model of male relationships. The low incidence of intragroup social interactions of any type in the focal males may reflect the energetic costs of social behavior. We suggest that intragroup social relationships among mantled howler males are structured by more subtle means than overt physical interactions, possibly including vocal communication, relationships with individual group females, and kinship.     

Sibships: Cooperation and competition among immature vervet monkeys

The relationships between siblings were compared with those between non-sibling peers amongst immature vervet monkeys (Cercopithecus aethiops) in three free-ranging social groups. Using the frequency and context of friendly, supportive, aggressive, and competitive interactions, sibships were found to stand out as associative units characterized by high levels of grooming and support during aggression. Competition between siblings was related to contests over access to maternal care, especially grooming, which may have been important in the establishment of relative sibling dominance. Siblings who were close in age tended to interact more frequently than those who were distant in age, but few sex differences were found. The close and supportive nature of sibling relationships appeared to affect the social structure of the groups.     

Interplay between various aspects in social relationships of young rhesus monkeys: Dominance,agonistic help,and affiliation

Relationships between group-living primates depend strongly on their position in the group dominance hierarchy and on their relationships with other group members. The influence of various behaviours on social relationships of immature rhesus monkeys (Macaca mulatta) was investigated here. Dominance ranks were established and related to the degree of affiliation in a dyad. Older monkeys were mostly dominant to the younger ones, regardless of kinship. Subordinate monkeys left proximity of their dominant members more often than they were left by them both among siblings and non-siblings, but the effect of dominance rank on the amount of play initiation and grooming in a dyad differed between these two types of dyads. The amount of agonistic help two individuals provided for each other was low among immatures. Nevertheless, pairs of siblings gave help to each other in agonistic conflicts more often than non-siblings, and such help was more often reciprocated between siblings than between non-siblings. Help in agonistic conflicts was positively correlated with the amount of time monkeys spent playing, grooming, or in proximity. Adults tended to interfere less in conflicts of frequent sibling play partners or non-sibling grooming partners. No evidence was found for young monkeys to exchange reciprocally grooming for agonistic help. It is argued that the time monkeys spend interacting with each other in affiliative interactions increases their familiarity and thus promotes close relationships between them. On the whole, young monkeys' relationships, like those between adults, are influenced strongly by their kinship, and position in the dominance hierarchy.     

A comparison of primiparous and multiparous mother-infant dyads inMacaca mulatta

The relationships of 42 rhesus monkey mother-infant dyads were examined to investigate the effects of the mothers' parity. Primiparous mothers were found to be younger, more excitable and less confident than the multiparous mothers. They also received more aggression from other adult females living in their social group and were more anxious about their infants, approaching and leaving them more frequently. The primiparous mothers protected their daughters more than the multiparous mothers and their daughters correspondingly spent more time in the ventro-ventral position and more time on the nipple. Their sons however spent less time in contact with their mothers than did the sons of the multiparous mothers. Although the primiparous mothers protected their daughters more than their sons, for the multiparous mothers the converse was true and sons received more maternal protection. It is suggested that the greater protectiveness of the multiparous mothers towards male infants may be a consequence of the interest siblings have in males, whereas the greater protectiveness of primiparous mothers to daughters may stem from their greater vulnerability to attacks from adult females. The importance of mother-mediated sibling influences may explain the lack of strong parity differences in previous studies.     

Social Relationships among Males in Multimale Siamang Groups

I quantified social and spatial interactions among adults in 4 multimale siamang (Symphalangus syndactylus) groups to evaluate the importance of aggression and avoidance in mediating male-male relationships. Actual genetic relationships among adults are unknown, but available mitochondrial data suggest that in 3 of 4 groups, neither male was the offspring or maternal sibling of the female, whereas in the fourth group, a matrilineal relationship between the female and 1 adult male was not excluded. Rates of aggression involving male-male dyads were very low. One male-female dyad maintained closer spatial cohesion than those of other adult dyads in 3 of 4 groups. Nonetheless, all adult males spent substantial percentages of their time ≤20 m from other adults in their groups. The percentages of time that male-male dyads spent in social grooming interactions did not differ from those of male-female dyads. In 3 groups, both males copulated with the group female. While previous studies have reported high rates of aggression between adult males and subadult male group members in siamangs, my results suggest that male-male relationships in multimale groups at Way Canguk were relatively harmonious. Acceptance of multimale grouping (and in some cases sexual polyandry) suggests that the benefits outweigh the costs under some circumstances. If there was a genetic relationship between males, then tolerance of delayed dispersal and copulation with the adult female may function as a form of parental investment. Males may also benefit from multimale grouping via enhanced territorial defense or reduced costs of mate defense.     

Breeding-season relationships of immature male rhesus monkeys with females. I. Individual differences and constraints on partner choice

Several aspects of breeding-season relationships with mature females are described for free-ranging immature male rhesus monkeys (Macaca mulatta)on Cayo Santiago. At puberty, immature males are still groomed by mature females, as they were as infants, but no longer receive active protection from females other than their mothers. A further, potentially beneficial, aspect of their associations with females lies in the opportunities to observe closely the tactical social and sexual interactions of receptive females with adult males. However, immature males themselves rarely copulate with females. Developmental constraints may explain the finding that most females observed by immature males were relatives but that high-ranking males showed a particular preference for adolescent relatives, whereas the preference of low-ranking males was for adult relatives. A further finding was that while there was a tendency for immature males to devote a greater proportion of their total observation time to females that ranked above their mothers than to those of inferior rank, these were also the females from which males received the most aggression, while most of the females with which the males groomed or copulated were lower ranking than the males’ mothers. Of three hypotheses concerning three possible types of social constraint which might account for such a finding, the data supported only one—that particular levels of aggression within these relationships are compatible only with certain other elements (observations), and not with others (grooming, copulation).     

Triadic interactions in Savanna-dwelling baboons

Interactions between adult males and infants may have important consequences not only for males and infants, but also for mothers. Considerable attention has been paid to interactions that involve two males and an infant. Investigators have proposed three different general strategies to account for this behavior: (a) exploitation of the infant for one male’s advantage, (b) protection of the infant from harassment and aggression, and (c) development of relationships with the infant’s mother. We review various accounts of these interactions, detail the hypotheses used to explain the behavior, make predictions derived from the hypotheses, and evaluate available data for testing the predictions. We conclude that multiple factors are probably at work, but the development of social relations between a male and an infant’s mother is a central force.     

Intergroup encounters in wild white-faced capuchins (Cebus capucinus)

Wrangham (1980) hypothesized that knowledge of the nature of intergroup encounters is crucial to understanding primate social relationships and social organization. I studied a single social group of wild white-faced capuchins over a period of 26 months and observed 44 encounters between social groups during 3703 hr of observation. All intergroup encounters consisted of predominantly hostile social interactions. However, nonaggressive interactions between males of different social groups occurred in a few cases. Adult males were the sole participants in 39 encounters and the primary participants in all 44 encounters. The alpha male was the most frequent participant. High-ranking females participated aggressively in five encounters, and low-ranking females never participated. There was no stable intergroup dominance hierarchy. I hypothesize that the need for male-male cooperation in intergroup aggression is an important factor influencing the quality of intragroup male-male relationships. Behavior during intergroup encounters is consistent with the idea that intergroup behavior is related to male reproductive strategies, but inconsistent with the idea that intergroup aggression is related to female defense of resources. The possibility that males are “hired guns” (Wrangham, 1980) cannot be ruled out.     

Social Relationships Among Male <Emphasis Type="Italic">Alouatta pigra</Emphasis>

We investigated male social relationships in 2 groups of black howlers (Alouatta pigra) in Palenque National Park, Mexico, over a 14-mo study characterized by frequent changes in male group membership. Both single males and pairs of males entered our focal groups. Single males tended to join groups, whereas pairs of males entering groups together successfully evicted resident males. The 19 male dyads across the 11 periods defined by changes in group membership were rarely in close proximity or interacting with one another. Nevertheless, 1 male formed significantly closer associations with resident females in 6 periods and achieved higher mating success during 4 of the periods when ≥1 female was sexually active. In the other 5 periods, no particular male maintained significantly closer associations with resident females, which may be a result of the rapid sequence of changes in male membership and corresponding social instability. Resident males participated frequently and consistently in mutual howling, but the resident male with the strongest female associations and highest mating success initiated howling bouts more frequently in all but 1 of the 6 socially stable periods. Though still preliminary, our findings suggest that variation in social relationships among male black howlers may be more related to their ability to establish strong relationships with females than to their relationships with one another.     

Mating patterns,mate choice,and birth season heterosexual relationships in free-ranging rhesus macaques

Birth season adult heterosexual nonkin relationships of 50 free-ranging female rhesus macaques (Macaca mulatta) in two social groups at Cayo Santiago, Puerto Rico were examined using focal follow (289 hr) and ad lib data. Eighty-eight percent of subjects had at least one relationship characterized by particularly high frequencies of spatial proximity, grooming, or both. These were designated “friendships.” Males intervened in aggressive interactions more frequently on behalf of Friends than non-Friends. Female aggressive support of males was extremely rare. Higher-ranking males experienced more friendships than lower-ranking males. High-ranking females had higher-ranking Friends than low-ranking females. Older females had higher-ranking Friends than younger females. Females groomed high-ranking Friends more than they were groomed by them, whereas they groomed low-ranking Friends less than they were groomed by them. In one social group, high-ranking females were more likely than low-ranking females to groom their Friends more than they were groomed by them. Males were more responsible than females for spatial proximity maintenance in 9 of 14 Friend dyads for which sufficient data were available. Neither male nor female dominance rank affected responsibility for proximity maintenance in Friend dyads. Eight of 24 females had friendships with males with whom they had completed copulations during their conception peri-ovulatory period of the preceding mating season. Two of 19 females completed peri-ovulatory copulations with Friends during the following mating season. Friendship was not correlated with either of two demonstrated female mate choice indicators: (1) proximity maintenance during estrus; or (2) cooperation with male “hip-grasp” courtship attempts. Males directed “muzzle-up” courtship signals at lower rates toward Friends than toward non-Friends. These and other investigators' results indicate that (1) protection from aggression is the primary benefit to female rhesus macaques of birth season heterosexual relationships; (2) the most effective protectors are in greatest demand as Friends; and (3) friendship has no effect or an inhibitory effect on mate choice in this species. Benefits to males of friendships were not apparent from this study but may include coalitional support against lower-ranking males.     

Sexual orientation, handedness, sex ratio and fetomaternal tolerance-rejection

Fraternal birth order (FBO) appears as a prenatal cause of 15% of homosexual males (gays) through mnemonic maternal anti-male factors. Non-right-handed men seem to be protected from homosexuality. Four hypotheses are proposed: (1) androgenic factors of non-right-handedness neutralize anti-male factors; (2) non-right-handedness and homosexuality are lethal or produce mental impairment; (3) non-right-handed male embryos are insensitive to anti-male factors; (4) mothers of non-right-handed fetuses do not produce anti-male factors. Studies of the sex ratio (SR) of older and younger siblings show: (1) a significant heterogeneity in the SR of siblings of right or non-right handed heterosexual men and women; (2) lesbians are born among siblings with high SR; (3) siblings of right-handed gays show a higher SR than non-right-handed gays that present a low SR. Based on our discovery of maternal tolerance-rejection processes, associated with genetic systems (ABO, Rh), where zygotes or embryos different from their mother induce better pregnancy and maternal tolerance than do those that share antigens with their mothers, I propose a new explanation for sexual relationships, sexual orientation, handedness and sibling SR. Lesbian embryos could induce tolerance from mothers with anti-female factors. Non-right-handedness could induce maternal tolerance, or change the maternal compatibility of "gay" embryos. Alternatively, gay embryos could be poor inducers of maternal tolerance towards male traits.