Male Migration in Barbary Macaques (Macaca sylvanus) at Affenberg Salem

We analyzed male migration during a 20-year period in the free-ranging Barbary macaque population of Affenberg Salem. Most natal migrations occurred around puberty, but only one third of all males left the natal group. Secondary group transfers were rare. All males immediately transferred to other bisexual groups. Migration rates were highest during periods with high adult female/male ratios within social groups. Immigrants highly preferred groups with fewer males of their own age than in the natal group, and many males immigrated into groups that had no male their own age. These groups originated from a skewed distribution of resident males during group fissions. A comparison of emigrants with their natal peers supports the inbreeding avoidance hypothesis as cause of emigration rather than the male competition avoidance hypothesis. Emigrants had no lower individual rank position and did not come from lower-ranking matrilines. Emigrants had more female maternal relatives, especially sisters. Males without female relatives almost never emigrated. Conversely, there is virtually no indication that emigrants were evicted from the natal group. Emigrants had no increased mortality. Paternity data revealed that the reproductive success of emigrants and natal males is similar, indicating that emigration had no reproductive cost. Many similarities between emigrants and natal males that separated from female maternal kin during group fissions suggest that inner migration during fissions is an alternative way to avoid maternal inbreeding. The mating system resulted in a genetic structure within social groups that largely diminished the chances for paternal inbreeding even without recognizing paternal kin.     

Inbreeding avoidance in rhesus macaques: Whose choice?

Whether nonhuman primates avoid copulating with close kin living in their social group is controversial. If sexual aversion to relatives occurs, it should be stronger in females than in males because of females' greater investment in each offspring and hence greater costs resulting from less viable offspring. Data presented here show that adult male rhesus macaques breeding in their natal groups at Cayo Santiago experienced high copulatory success, but copulated less with females of their own matrilineages than with females of other matrilineages. Adult females were never observed to copulate with males of their own matrilineage during their fertile periods. Although natal males sometimes courted their relatives, examination of two measures of female mate choice showed that females chose unrelated natal males over male kin. Female aversion to male kin was specific to the sexual context; during the birth season, females did not discriminate against their male relatives in distributing grooming. Evolved inbreeding avoidance mechanisms probably produce different outcomes at Cayo Santiago than in wild rhesus macaque populations. Gender differences in sexual aversion to relatives may be partly responsible for differences between studies in reported frequency of copulations by related pairs. © 1993 Wiley-Liss, Inc.     

Group fission in a semifree-ranging population of Barbary macaques (Macaca sylvanus)

During a 16-month study of semifree-ranging Barbary macaques (Macaca sylvanus) the group under observation divided into two groups. Observations were carried out in 1987–1988, at «La Montagne des Singes,” Kintzheim, France. A subgroup of monkeys, which was already cohesive at the beginning of the study, became progressively autonomous in relation to the rest of the main group, during the mating season. Overt aggression between the males of the two groups during this period brought about the fission. Only low-ranking genealogies left their group of origin. Dominance relations between females remained identical in both groups except for one lineage. The alpha male and the alpha female of the subgroup had a close relationship before the fission occurred. The sequence of agonistic intergroup relations is described and analyzed in relation to male sexual competition and female alliance power. The results suggest that: (1) the males of the subgroup instigated the fission because it was the best strategy for them to counter sexual competition; and (2) the females followed the males in order to maintain their alliance network, necessary to insure their dominance status over subordinate females.     

Dispersal patterns in Costa Rican mantled howling monkeys

Both male and female juveniles disperse in Costa Rican mantled howling monkeys (Alouatta palliata). 79% of the males and 96% of the females leave their natal groups. Males may spend up to 4 years and females up to 1 year as solitaries. Extra-group individuals are faced with only three possibilities, i.e., form a new group by joining another extra-group individual, join an established social group, or remain solitary. Most surviving extra-group individuals join an established social group which contains no kin. Females join with the help of a resident male and once in a group proceed to rise to the alpha position through dyadic interactions. The immigrant female either becomes the alpha female or leaves and tries again in another group. Males challenge the alpha male and either defeat him or remain solitary. Competition with relatives for limited high quality food may be the reason for both sexes leaving their natal groups in howlers. By leaving, the successful immigrants increase their mothers inclusive fitness while suppressing the fitness of nonrelatives instead of remaining natal and competing with relatives for limited food.     

Group fission in Nilgiri langurs (Presbytis johnii)

During a field study on habituated groups of wild Nilgiri langurs (Presbytis johnii),four cases of group fission were observed which differed markedly from group changes reported in other species of the same genus. All fissions occurred in groups with more than one mature male and roughly coincided with the onset of loud call vocalization by the second mature male group member. As a result of the fission the founder group factions acquired the structure of a onemale group. The filial factions consisted of both mature males and females and occupied, at least temporarily, a part of, or an area adjacent to, the founder group. During fission, the amount of aggressive interactions between the two mature males increased. However, the majority of agonistic interactions involved ritualized threat and intimidation behavior without physical aggression. In at least three cases the two mature males involved in the fission had not joined recently but had lived in these groups for three years or more. Although the presence of all male bands and single males in the two study populations indicate that male replacement by invasive takeovers may occur, it is suggested that in Nilgiri langurs, noninvasive fissions are probably another common mechanism for the foundation and maintenance of bisexual one-male groups.     

Postdispersal nepotism in male long‐tailed macaques (Macaca fascicularis)

Cooperative behaviors are promoted by kin selection if the costs to the actor are smaller than the fitness benefits to the recipient, weighted by the coefficient of relatedness. In primates, cooperation occurs primarily among female dyads. Due to male dispersal before sexual maturity in many primate species, however, it is unknown whether there are sufficient opportunities for selective tolerance and occasional coalitionary support for kin selection to favor male nepotistic support. We studied the effect of the presence of male kin on correlates of male reproductive success (residence time, duration of high dominance rank) in non‐natal male long‐tailed macaques (Macaca fascicularis). We found that “related” (i.e., related at the half‐sibling level or higher) males in a group have a significantly higher probability to remain in the non‐natal group compared to males without relatives. Moreover, males stayed longer in a group when a relative was present at group entry or joined the same group within 3 months upon arrival. Males with co‐residing relatives also maintained a high rank for longer than those without. To our knowledge, this is the first demonstration of a potential nepotistic effect on residence and rank maintenance among non‐natal males in a social system without long‐term alliances.     

Intergroup transfer and incest avoidance in semifree-ranging barbary macaques (Macaca sylvanus) at Salem (FRG)

During a long-term study of social behavior of semifree-ranging Barbary macaques, data on group transfer and sexual behavior were collected. A large population containing five groups living in a 14.5 ha outdoor enclosure was studied. Demographic data on the whole population are available for a period of six years. Data on sexual behavior of members of one large group were collected, by both focal animal and ad libitum techniques, for a period of two years. Matrilineal kinship relations of all except the oldest members of this intensively studied group were known. Males changed groups most frequently between the ages of three and five years i.e., the time at which sexual maturity was reached. While more than 30% of all males three years old and older changed groups during the study period, many males remained members of their natal group even after reaching adulthood. However, sexual interactions, especially matings between close matrilineal relatives, were found to be almost absent, suggesting that Barbary macaques are not highly inbred as formerly supposed.     

Dynamics of fission in a wild Barbary macaque group (Macaca sylvanus)

We studied the dynamics of a group of Barbary macaques (Macaca sylvanus) in Algeria from March 1983 to November 1989. Troop fission began in autumn 1987, when group size had more than doubled, to include 76 animals. We observed 11 temporary splits of this group during the mating seasons of 1987 and 1988. The process was interrupted during the 1988 birth season. In June 1989, fission resumed and ended with the formation of three independent groups that included 50, 24, and 13 individuals. Adult females played an important role in the process of fission. They initiated the rapid formation of two, and later three, coherent nuclei, distributed in two or three bisexual subgroups; on several occasions these nuclei also formed subgroups without any adult males. Adult males remained together in a single nucleus for longer periods of time than females did. However, during fission, 35% (N=20) of resident males emigrated to neighboring groups, while 11 strange males immigrated into the focal group; over 6 years, 57% of the male transfers occurred after the beginning of the process. After group fission, maternally related individuals lived together in the new groups. The majority of resident males remained with the largest of the three groups, while most of the immigrant males were in another group. The third group included a single adult male. Possible factors that induce group fission are discussed.     

Origin and fate of a one-male savanna babbon group formed by fissioning

An adult male chacma baboon,Papio hamadryas ursinus, emigrated from his habituated natal group at age 100 months and joined a neighboring multimale-multifemale group. Subsequently this male, together with eight adult females, all with infants <1 year old, formed a one-male group (OMU). The male was the only adult male in this group for over 200 days. Circumstantial evidence shows that shared parentage of the infants with this male determined which females joined the new group. In addition, 10 juveniles younger than 4.5 years joined the new group. Based upon persistent close physical association, especially at sleeping sites, these juveniles were presumed to be the older independent progeny of the eight adult females. Thus, the founding elements of the new group were the adult male and partial matrilines—mothers, their infants, and their independent juvenile offspring. The OMU status of the group ended with the sequential intrusion of three males known to be low ranking in other groups. The first of these intruders was successful in joining the OMU in spite of the OMU male's earlier successes in rebuffing this and other potentially high-ranking males. Six additional males later entered the fission group during a 6-month interval. Two of them outranked the original male. All additional males present in the last 100 days of observations came from the same group, which was not the group of origin of the females.     

Age-related and individual differences of reproductive success in male and female barbary macaques (Macaca sylvanus)

Age-related and individual differences in longterm reproductive success were analyzed in two social groups of free-ranging Barbary macaques. Maternity data were obtained from continuous birth records and paternity was determined with oligonucleotide-fingerprinting. The fathers of 246 of 286 investigated individuals could be identified. They were born during a 14-year period and represented 73 and 34% of all known offspring from the females of the study groups B/F and C, respectively. Only these infants were considered when comparing male reproductive success with that of females. The necessary adjustment of the female data resulted in small deviations from the true values in one group, but substantially increased individual differences in female fertility in the second group. Subadult males, 4.5 – 6.5 yrs old, had a much lower reproductive success than adult males (7.5 – 25 yrs old) and same-aged females. Reproductive success of adult males was not significantly affected by age, while females invariably ceased reproduction during the first half of the third decade of life. Males were more likely than females to leave no offspring, unless they survived 9 – 10 yrs of age. The number of years with breeding opportunities was important for male reproductive success but less significant than that for females. Reproductive success of several males during the 14-year study period was similar to or even exceeded that possible for a female in her whole lifetime. Variance of male reproductive success significantly exceeded that of females in both study groups.     

Group Fission in Moor Macaques (Macaca maurus)

This is the first report of group fission in a wild group of Moor macaques (Macaca maurus) in South Sulawesi, Indonesia. The subject group, which has been observed on the basis of individual identification since 1988, showed no sign of fission in April 1999. In August 1999, the group had split in two, with the same number of mature females in each new group. For the most part, mothers and their offspring joined the same groups. Dominance relations and association patterns established during the previous year among adult females did not strongly affect new group membership. The difference in female reproductive state between the two branch groups was a prominent characteristic. The -male of the original group visited both groups at the first stage of group fission, even though otherwise the compositions of the new groups were stable. After the division, six adult males from outside the original group immigrated exclusively into the group that did not contain the -male of the original group. Severe intergroup encounters occurred between the two groups. We discuss the process of the fission and the resultant pattern in relation to the egalitarian dominance style among females, lack of seasonality in reproduction, and resemblance to one-male type social organization.     

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.     

The Influence of Kinship on Familiar Natal Migrant Rhesus Macaques (Macaca mulatta)

In most primate species, females remain in the natal group with kin while males disperse away from kin around the time of puberty. Philopatric females bias their social behavior toward familiar maternal and paternal kin in several species, but little is known about kin bias in the dispersing sex. Male dispersal is likely to be costly because males encounter an increased risk of predation and death, which might be reduced by dispersing together with kin and/or familiar males (individuals that were born and grew up in same natal group) or into a group containing kin and/or familiar males. Here we studied the influence of kinship on familiar natal migrant rhesus macaques (Macaca mulatta) on Cayo Santiago, Puerto Rico, by combining demographic, behavioral, and genetic data. Our data suggest that kinship influences spatial proximity between recent natal immigrants and males familiar to them. Immigrants were significantly nearer to more closely related familiar males than to more distantly related individuals. Within a familiar subgroup, natal migrants were significantly closer to maternal kin, followed by paternal kin, then non-kin, and finally to males related via both the maternal and paternal line. Spatial proximity between natal immigrants and familiar males did not decrease over time in the new group, suggesting that there is no decline in associations between these individuals within the first months of immigration. Overall, our results might indicate that kinship is important for the dispersing sex, at least during natal dispersal when kin are still available.     

The fitness consequences of kin-biased dispersal in a cooperatively breeding bird

Cooperative alliances among kin may not only lead to indirect fitness benefits for group-living species, but can also provide direct benefits through access to mates or higher social rank. However, the immigrant sex in most species loses any potential benefits of living with kin unless immigrants disperse together or recruit relatives into the group in subsequent years. To look for evidence of small subgroups of related immigrants within social groups (kin substructure), we used microsatellites to assess relatedness between immigrant females of the cooperatively breeding superb starling, Lamprotornis superbus. We determined how timing of immigration led to kin subgroup formation and if being part of one influenced female fitness. Although mean relatedness in groups was higher for males than females, 26% of immigrant females were part of a kin subgroup with a sister. These immigrant sibships formed through kin recruitment across years more often than through coalitions immigrating together in the same year. Furthermore, females were more likely to breed when part of a kin subgroup than when alone, suggesting that female siblings form alliances that may positively influence their fitness. Ultimately, kin substructure should be considered when determining the role of relatedness in the evolution of animal societies.     

Limited indirect fitness benefits of male group membership in a lekking species

In group living species, individuals may gain the indirect fitness benefits characterizing kin selection when groups contain close relatives. However, tests of kin selection have primarily focused on cooperatively breeding and eusocial species, whereas its importance in other forms of group living remains to be fully understood. Lekking is a form of grouping where males display on small aggregated territories, which females then visit to mate. As females prefer larger aggregations, territorial males might gain indirect fitness benefits if their presence increases the fitness of close relatives. Previous studies have tested specific predictions of kin selection models using measures such as group‐level relatedness. However, a full understanding of the contribution of kin selection in the evolution of group living requires estimating individuals' indirect fitness benefits across multiple sites and years. Using behavioural and genetic data from the black grouse (Tetrao tetrix), we show that the indirect fitness benefits of group membership were very small because newcomers joined leks containing few close relatives who had limited mating success. Males' indirect fitness benefits were higher in yearlings during increasing population density but marginally changed the variation in male mating success. Kin selection acting through increasing group size is therefore unlikely to contribute substantially to the evolution and maintenance of lekking in this black grouse population.     

Consequences of group fission for the patterns of relatedness among rhesus macaques

When mammalian social groups exceed their optimal size, they often tend to split. In view of the potential evolutionary benefits, it should be more advantageous for animals to stay with kin, rather than nonkin, during such fission events. In the present study, the spontaneous fission of two social groups, R and S, of rhesus macaques living on Cayo Santiago, Puerto Rico, provided the opportunity to compare the kinship structure of the corresponding parent and daughter groups, using information on both maternal and paternal relatedness. In both instances, maternal half-siblings and pairs of animals from the same family were significantly more prevalent in the fission products than in the parent group. During the split of group R, significantly more paternal half-siblings stayed in the remnants of the parent group than joined the seceding group. Our findings are compatible with previous behavioural studies demonstrating that female primates bias their social behaviour more to maternal than to paternal kin, but that both types of half-siblings prefer each other more than unrelated animals. It remains to be clarified by future research, however, whether the observed co-segregation of paternal half-sibs in our study reflects active choice or is a by-product of the group-specific kin structures, prior to fission.     

Natal emigration by both sexes in the La Pacifica population of mantled howlers: when do some stay?

We have reported previously that all male and female mantled howlers emigrate from natal groups at Hacienda La Pacifica, Costa Rica. In the years since that report, a small number of juveniles have stayed in the natal group without experiencing a solitary phase. Here, we present a post hoc analysis on juvenile emigration in six groups of howlers under observation for varying amounts of time between 1972 and 2005. Our records revealed 139 juveniles for whom emigration status was certain, and 125 of these did emigrate. There was a significant association between presence of mother and emigration: juveniles without mothers were more likely to remain in their natal group (chi(1)(2) = 53.1, P<.0001). The mean age of emigration for all juveniles (n = 125) was 2.47 years (SD = 0.9, range = 1.5-6.5). There was no difference in age of emigration by adult male composition (one-male, multi-male, both), but juveniles of unknown sex emigrated younger than either known males or females (F(2,116) = 4.4, P<.02). For emigrating juveniles of known sex (n = 99), both males and females without mothers left at a later age than those with mothers (F(1,95) = 6.5, P<.02). Although philopatry or delayed emigration occurs in a few motherless animals, most males and females do emigrate from their natal groups at ages consistent with those reported for other species of howlers.     

Group fission among wild toque macaques as a consequence of female resource competition and environmental stress

At Polonnaruwa, Sri Lanka, four out of 29, groups of toque macaques, Macaca sinica, divided in a period of 16 years. Temporary peripheral subgroups of varying sizes and compositions preceded fission by 9–40 months. Fission crystallized within a month through an increase and stabilization of subgroup membership and permanent division. All members in the newly seceded groups had been frequent participants in pre-fission subgroups, and belonged to subordinate matrilineages. Subgroups, and hence group divisions, were initiated by cores of mutually loyal females and occurred mostly along kinship lines. In the year of fissions, the rate of change in female dominance relations was significantly greater among groups that divided than among those that did not. It is hypothesized that low-ranking females secede to form new groups when the costs, especially of intragroup competition for food resources, outweigh the benefits of group membership. Such seceding females were easily available and familiar mates for group males that had recently lost rank. Final division, therefore, resulted from a coalition of subordinate females and males acting according to their respective interests. It was triggered in this population by rapid growth of some groups to large size and by environmental stress (the reduction and fragmentation of food resources caused by drought and a cyclone), which accentuated the costs of resource competition. Male aggression, such as infanticide, which negatively affects female fitness, might also have contributed to one group fission.     

Socio-sexual factors of troop fission in wild Japanese monkeys (Macaca fuscata yakui) on Yakushima Island,Japan

The troop fissions which occurred in a wild population of Japanese monkeys (Macaca fuscata yakui) were observed from 1977 to 1979 on Yakushima Island. The fissions were initiated in the breeding season by non-troop males who established a consort relation with estrous females. In order to analyze the socio-sexual factors which accelerated the fissions, the male emigrations and immigrations before and after two successive fissions, and the copulation frequencies, competition among males and preferences of mating partners in both sexes in the 1977–78 breeding season after the first fission were examined. The results indicated that three factors (a large number of non-troop males, a shortage of troop males and the females' choice of mating partners) effectively influenced on the establishment of consort relationships between non-troop males and estrous females. It is suggested that these factors may exert different effects on the troop disorganization in relation to troop size. In small-sized troops, a large number of non-troop males and a shortage of troop males may lead to stronger competition between them, and the females' choice affected by prolonged intimate relations with the dominant TMs may reduce their priority of access to estrous females. This situation possibly stimulates fission or male emigration in small-sized troops under the natural conditions on Yakushima Island. In contrast, in large-sized troops under isolated conditions, a surplus rather than a shortage of troop males may contribute to troop disorganization, as most former studies have suggested. A higher socionomic sex ratio may decrease the mating activities of subordinate troop males and increase the competition among them. This situation possibly accelerates the fission of large-sized troops through prolonged interactions between females and subordinate or peripheral troop males. A lower ratio and the females' choice, however, raise the mating chances of subordinate troop males and may not promote the fission of large-sized troops under isolated conditions. This study was financed in part by a Grant-in-Aid for Special Project Research on Biological Aspects of Optimal Strategy and Social Structure from the Japan Ministry of Education, Science and Culture, and by the Cooperative Research Fund of the Primate Research Institute, Kyoto University.     

Population and family studies on Kuru

Abstract

A model of group fissioning, applied by Mange (1964) to a Hutterite population and by Chagnon (1975) to an Amazonian Indian population, is applied to the “Nebraska” Amish community in Central Pennsylvania. Genealogical data were collected on 228 living, married individuals and their known ancestors in two new churches resulting from the fission of a church in 1978. The acrimonious fissioning of the church produced increased levels of intrachurch genealogical relatedness in the two new churches formed, as compared to the level of relatedness in the original church, with a greater increase for male/male relatedness than for female/female relatedness. This general increase in relatedness, documented for three human populations and also for fissioning groups of rhesus macaques, conforms to predictions from kin selection theory.