Social interactions among adult male langurs (Presbytis entellus) at Rajaji Wildlife Sanctuary

A population of langurs (Presbytis entellus)at the Rajaji Wildlife Sanctuary in northern India was investigated for 1820 hr throughout a 10-month period in 1978. Data were collected from four bisexual troops and the adult males that ranged outside of bisexual troops. Most (60%) of the observation hours occurred with a main study troop from which social and ecological data were collected. The langur population at Rajaji shows pronounced birth and mating seasons. The population density is high (ca. 80/km ²), with about 75% of the adult males living outside of bisexual troops, which typically are large and multimale. Males outside of bisexual troops occur in small all-male bands or as isolates. Relations between bisexual troops and all-male bands are characterized by relatively low levels of aggression, and members of all-male bands are able to associate with bisexual troops for prolonged periods during the mating season. As a result of these associations, nontroop males are about as successful as troop males in achieving reproductive access to troop females. These associations between bisexual troops and all-male bands occurred with a minimal amount of agonistic behavior and without mortality or injury to troop females or immatures.     

Social Relationships Among Ring-Tailed Lemurs (Lemur catta) in Two Free-Ranging Troops at Berenty Reserve, Madagascar

We observed two free-ranging troops of ring-tailed lemurs at the Berenty Reserve, Madagascar. Kinship affinities in these troops are known only for mothers and their offspring 4 years of age. We attempted to quantify social relationships. Almost all agonistic interactions were dyadic, and triadic agonistic interactions, such as alliances, were very rare. Dominance hierarchies in both sexes in the two troops were not linear. As in cercopithecine monkeys, mothers were dominant over their adult daughters. However, the daughters were not ranked immediately below their mothers. Close proximity and social grooming occurred more frequently between closely related females, such as mother–daughter and sister–sister dyads, than between unrelated females. Frequent-proximity relations also occurred between adult males that had emigrated from another troop and entered the present troop together, even though they did not rank closely to one another. Subordinates were likely to groom and to greet dominants more frequently than vice versa. During group encounters, particular females were involved in agonistic interactions with animals of other troops, regardless of dominance rank. Adult males, regardless of their dominance rank, but not adult females, constantly tried to drive solitary males away.     

Symmetrical patterns in non-agonistic social interactions found in unprovisioned Japanese macaques

Non-agonistic social interactions in an unprovisioned troop of Japanese macaques (Macaca fuscata yakui) were analyzed with the spacing between individuals, leading-following interactions, and exchange of social grooming. The most frequent interactions were found between kin-related females. Unrelated females stayed with one another rather frequently, but rarely exchanged social behaviors. Interactions between males and females were infrequent though they were occasionaly observed between high-ranking males and high-ranking females. Very frequent exchange of grooming was observed between males, and even high-ranking males exchanged grooming more frequently with males than with females. Most non-agonistic social interactions in the study troop were based on bidirectional exchange of social behaviors, in which no clear tendency relevant to dominance or sex was found; while in provisioned Japanese macaque troops, associations between males and females, between unrelated females, and between males were formed mainly be subordinates' active roles in associative behaviors. This seems relevant to the idea that dominance grealty influence social life in provisioned troops. The present study provides guidelines for interspecific comparison of social interaction patterns of macaque species.     

Feeding and ranging patterns of forest hanuman langurs (Presbytis entellus)

The feeding and ranging patterns of a troop of hanuman langurs (Presbytis entellus, Colobinae) were studied in Kanha Tiger Reserve, central Indian Highlands for 1850 hr (1981–1982), in a mosaic of moist deciduous forest and anthropogenic meadow. The location, size, and species of each tree within the 74.5-ha troop annual range was known and the phenology of all tree species was sampled. According to scan sampling, the troop spent 25.7% of the daytime feeding, with range use concentrated on an island of dry deciduous forest. Whereas adjacent troops occupied only the periphery of the focal troop's range, all-male bands occupied its center, especially during takeover and infanticidal attacks. The troop consumed items from 60 of the 67 species of trees and woody climbers available; mature leaves (34.9% of feeding time), fruits (24.4%), leaf buds (10.6%), flowers and flower buds (9.5%), young leaves (3.6%), insects (3.0%), and gum (1%). The monthly utilization of fruit, open leaf buds, and flower buds is correlated significantly with their abundance, and the troop spent significantly more time feeding and less time moving when consuming mature leaves. Comparison of tree dispersion and langur ranging patterns suggests that the distribution of the most important food trees is a major influence on their range use.     

Changes in dominance rank and division of a wild Japanese monkey troop in Arashiyama

This article forms the second report on the Arashiyama troop of Japanese monkeys and concerns a troop division which took place in June, 1966, and various problems of rank and consanguinity which accelerated the division. (1) The hypothesis advanced in the first report has been verified; (2) at the time of troop division, several consanguineal groups formed one unit; (3) among 16 consanguineal groups, those from 1st to 7th in rank joined the A troop, while those from 8th to 16th joined the B troop; (4) dominance relation between the two division troops was B troop>A troop, reflecting the former ranking between the leader males of the two troops; (5) shifting of monkeys from one troop to the other after division occurred frequently, but males began to make their own movements when they attained 4 or 5 years of age and rarely moved together with their mothers or other consanguineous-relatives; (6) monkeys which were continuously in the same troop after division almost always obtained higher ranks than did monkeys who frequently shifted from one troop to the other; (7) after division, some males joined neither of the two division troops but formed a group, a so-called all-male group or male party, and moved about independently.     

Population organization of wild pig-tailed macaques (Macaca nemestrina nemestrina) in West Sumatra

A field study on wild pig-tailed macaques was conducted in West Sumatra, Indonesia, during three periods from January 1985 to February 1987. During the nine months of the first two periods, unprovisioned monkeys were traced and observed. During the eight months of the last period, monkeys were provisioned and observed mainly at baiting sites. Three troops and ten solitary males appeared at the two baiting sites. Some males immigrated into and emigrated from the troops. The troops had a multi-male multi-female composition. The size of the various troops was 74, 49, and 81 individuals, respectively, and the mean adult sex ratio in the troops was 1:6.3; that is, markedly biased towards females. The home ranges of two of the troops overlapped considerably. When the troops encountered each other at the baiting sites, a clear dominance relationship was recognized. The troops differed in their integration as ranging units: two of the troops did not form subgroups (temporary fission and fusion of each troop), while the other troop frequently split into subgroups. Recent field studies on pig-tailed macaques have suggested a multi-leveled society with harem-type unit groups. However, in the present study, the troops observed had neither a substructure similar to harem-type groups nor a superstructure that emerged as a result of fusion of the troops. The unit group of the pig-tailed macaques appears to be a multi-male, matrilineal group.     

Ecology and behavior ofPresbytis thomasi in Northern Sumatra

A field study of 23 bisexual troops, ranging in size from 3 to 21 members ( =8), and two all-male groups of the Thomas's leaf monkey (Presbytis thomasi) was conducted in North Sumatra from November 1981 to April 1984. Most troops (N=19 or 82.6%) contained only one adult male. Two troops and one group were most intensively studied. The home range was 12.3–15.7 ha for the two bisexual troops, and only 1.7 ha for the one, all-male, ten-member group. Fruits composed more than 50% of their diets. Vocalizations were classified into 13 types. Births occurred at any time of the year. Among three males of a bisexual troop, serious fights were observed: two males died of wounds and the former beta male became the new alpha male. After this social change, the home range area of this troop gradually shifted eastward. But, 3.5 months after the social change, a 9-month-old infant male stayed alone in the western part of its former range. Thereafter, he became a solitary male and sometimes went into the riverine area of the Bohorok river. The occurrence of male replacement suggests instability of multi-male organization in bisexual troops. Moreover, the different mortality rate between males and females and the unequal sex ratios forced by the formation of one-male troops, maintained high tension levels among males competing on females.     

Inter-male associations in a wild Japanese macaque troop on Yakushima Island,Japan

Adult male association and its annual change were studied in a wild population of Japanese macaques (Macaca fuscata yakui) on Yakushima Island, Japan. Unlike many other Japanese macaque troops, adult troop males frequently maintained proximity and exchanged grooming with one another in both the mating and non-mating seasons, and the dominance relationship rarely appeared in such inter-male associations. The few cases of agonistic interactions occurred mostly when estrous females or food resources were immediately concerned. Although troop males were very intolerant to newly appeared solitary males (new males) during the mating season, close associations were formed between troop males and new males as soon as the mating season terminated. The consort of new males and lower-ranking troop males with estrous females was frequently disturbed, but these males could copulate no less frequently than higher-ranking males. A comparison among macaque species suggests the existence of two forms of inter-male association: (1) the frequent association based on the symmetrical exchange of social behaviors; and (2) the infrequent and asymmetrical association related to the dominance relationship. The form of inter-male association seems to be influenced by whether or not males can keep close associations with females throughout the year.     

Group size variability in primates

The effects of mean troop size, diet, territoriality, and habitat upon temporal variability of group size in primates were investigated using variance functions relating mean group size and temporal variability. Two different types of variability were described: (1) within group variability where a single troop was followed over a given period of time; and (2) between group variability where the author(s) did not distinguish one troop from the other. In the second category, CV (SD/mean) as an index of temporal group size variability proved to be dependent on mean group size among the Cercopithecidae. Large groups are more unstable in size than small ones. In the Cebidae, variability was independent of mean group size and therefore large groups are as variable through time as small ones. Ecological factors showed no effects on the observed level of between group variability. Within group variability was found to be smaller than the level of between group variability in all species tested. The results are related to social organization and to the degree of feeding interference observed within and between troops. Future practical applications for our results are considered.     

An ecological study of troop fissions of Japanese monkeys (Macaca fuscata yakui) on Yakushima Island,Japan

Wild, habituated, Japanese monkeys were observed from 1975 to 1979 on Yakushima Island, Southern Japan. The monkey troops had a continuous distribution in a warm temperate forest. Demographic data on local populations was collected. The population density was 33 animals/km². The growth rate of the studied troop was 3.0% per year. A significant correlation between home range areas (R) and troop size (P) was found (r=0.955,p<0.005), using anR-P equation,R=1.84P. One troop split into three troops through two successive fissions. Twenty-one intertroop encounters were observed. Five types of encounters were distinguished. The encounters were apparently territorial defence. Increases in birth rate and socionomic sex ratio after the fissions were prominent. The following four factors had a direct effect upon the dispersion of the troops after fission: (1) dominance relation between the fission troops; (2) social pressure of the neighbors; (3) troop's attachment to its home range; and (4) structure of the environment. The home range of Japanese monkeys is a territory, and territoriality is a population regulating mechanism which serves to reduce competition for food.     

Intertroop Transfer and Dominance Rank Structure of Nonnatal Male Japanese Macaques in Yakushima, Japan

We examined the interaction between intertroop transfer and male dominance ranks in a wild population of Japanese macaques (Macaca fuscata yakui) in Yakushima using data collected over 15 years. Intertroop transfer tended to maintain a linear, stable, and age-graded dominance rank order among nonnatal males irrespective of variation in troop size or composition. All males that joined a troop at the top of the rank order were prime adults. Among males joining at lower ranks, entry at the most subordinate position in the hierarchy was common. Males joining at lower ranks tended to join troops in which all other resident males were the same age or older. Adult males tended to join troops with few or no males. Young males tended to join troops with many resident males, and in which a relatively large proportion of males was other young ones. Intertroop transfer was responsible for most rank changes of resident males. The most common cause of males rising in rank was the emigration or death of a higher-ranking male. Males fell in rank most frequently as a result of a new male joining the troop at the top of the hierarchy. Rank reversals among resident males were rare. The cumulative effects of male transfers produce sociodemographic variation within a troop over time and sociodemographic diversity among troops in a local population. A key feature of intertroop diversity is that larger troops have a significantly greater proportion of young males than smaller troops. This diversity also creates the potential for intertroop variation in the severity of male competition and provides a range of options for transferring males.     

The role of play in social organization: Comparative observations on squirrel monkeys (Saimiri)

Various functional theories of play stress that social play is essential for the practice and learning of sex roles, dominance relationships, troop culture, integration of individuals into the troop structure, the control of aggression, etc. Data on squirrel monkeys (Saimiri) in natural environments indicate that social interaction and troop integration can develop in various manners in the absence of social play.Comparative observations were made on squirrel monkeys in a seminatural environment in Florida and 43 natural environments in Panama, Colombia, Peru, and Brazil. There was a broad range of variance in the data on ecology, troop size, troop cohesiveness, average individual distances, frequency of play, etc. In some environments, individuals in the infant and juvenile age classes engaged in social play for approximately 1.5 to 3 hours a day. However, in one environment, not a single incidence of social play occurred during 261 hours of close range observation. The troops in which no play occurred were very cohesive (i.e., they seldom fragmented), and the animals traveled at close individual distances. Agonistic interactions were not uncontrolled. Copulations were observed; and 85 percent of the adult females were accompanied by infants, which indicates a normal rate of reproductive success for the species.Data are presented on friendly, aggressive, sexual, and spacing behavior in squirrel monkeys. These data indicate that (1) social play is not necessary for the development and/or learning of an adaptive modicum of social interaction patterns and troop cohesion, but (2) the opportunity to play provides learning experiences in which young animals can develop more complex, varied social interaction patterns and stronger habits for engaging in frequent social exchanges.     

Comparative study of grooming relationships among wild Japanese macaques in Kinkazan A troop and Yakushima M troop

The influences of socionomic sex ratio (SSR; adult males/adult female) and troop size upon male-male, female-female, and male-female grooming relationships were examined and compared between two wild Japanese macaque troops (Kinkazan A and Yakushima M troops) in Japan. The Yakushima M troop was smaller and had a higher-SSR than the Kinkazan A troop. Between the troops, (1) the male-male grooming frequency and number of partners were greater in the Yakushima M troop than in the Kinkazan A troop; (2) the female-female grooming frequency and number of partners were not different; and (3) the male-female grooming frequency and number of partners were not different. Based on these features, the patterns of female-female and male-female grooming relationships appear to be independent of SSR and troop size variations. In contrast, male-male grooming relationships are influenced by both factors, especially SSR. Frequent grooming interactions among males may be useful for the continued coexistence of relatively many males especially in a higher-SSR troop.     

Intermale relations and troop male membership changes in langurs (Presbytis entellus) in Nepal

Data on intermale social relations and troop membership changes in one Nepalese high-altitude population of free-ranging langurs (Presbytis entellus)are reported here. Data were collected from six troops by three observers and cover 32 months of observations. The predominantly multi-male troops indicate an alternating pattern of exclusions and introductions with gradual adult male replacement. Takeovers and infant killing were not observed. Analysis of adult social behavior records show qualitative and quantitative differences in intrasexual relations, with primarily agonistic social contacts occurring between males. Agonistic encounters between females and between males differ in frequency of occurrence, types of be-haviors used, cause, and consistency in direction of threats between individuals. Individual adult male frequency of interaction with females and immatures varied significantly, with the majority of these interactions occurring between the dominant troop male and other troop members. Data indicate that intermale dominance is a major factor in determining male access to fertile females: This appears to be achieved by either directly excluding males from the troop or effectively “controlling” their inter-actions with troop females. Data from these studies are compared with data from other Presbytis entellusinvestigations. Review of these data suggests that intraspecific variability in intermale social dynamics and type of troop male membership change are correlated with the percentage of nontroop males. It is suggested that environmental pressures resulting in social crowding can be critical in determing the occurrence of takeovers in some populations of Presbytis entellus.     

Intertroop relationships among Japanese monkeys

Intertroop relationships among Japanese monkeys, which have been paid only scant attention for the past 20 years, are examined from several points of view. Japanese monkey troops are generally distributed in such a way as to concentrate locally, that is, to form a local concentration of troops (LCT). About 20% of the nomadic ranges of the troops within LCT's overlap; but in their natural state, they seldom approach but rather to avoid one another. From observations of intertroop encounters at Takasakiyama, where three troops are provisionized and use the same feeding place, it has been found that there exists a dominant-subordinate relationship among troops, that monkeys of each troop have a clear consciousness of belonging to their troop, and that monkeys of different troops rate one another on the basis of their capability. The frequency of male transfer between troops within LCT's is by far higher than between LCT's. In Japanese monkey society, a troop only is a social unit and a social order higher than a troop is not seen; however, it is not impossible to consider an LCT a consanguineally connected group by reason of the transfer of males among the troops within it.     

Sociological study on the troop fission of wild

During the period from June to July 1983, the Hanyama-A troop of wild non-provisioned Japanese monkeys on Yakushima Island began to show signs of troop fission. Adult females together with their infants and juveniles subdivided into two groups, the Hanyama-K group and Hanyama-M group. After the subdivision, all of the troop males were observed vacillating between these two female groups. During the mating season, non-troop males were also observed moving around the two female groups. After this mating season, one of these non-troop males was found to have entered and become the alpha male in one of the groups, while higher-ranking adult males of the original troop settled into the other group. Each fissioned group was strongly considered to be composed of either high-ranking matrilines or low-ranking matrilines as observed previously in provisioned troops. The dominance relation between the two fissioned groups indicated that dominance rank reversal between these two female kin groups must have occurred during the course of subdivision of the troop. However, different from most previous cases of troop fission, there was no indication that males ever participated in the subdivision of the original female group. This was disrupted not as a result of males' involvement, but only as a result of antagonism among females, which initiated the troop fission. The main factor which appeared to determine when and in which fission group males eventually settled was the competition between the troop males' coalition and non-troop males and their ability to monopolize females. The present process of troop fission suggests a dual strategy between males and females (Wrangham, 1979, 1980) even in the society of Japanese macaques.     

Better to Live with Allogenerics Than to Live Alone? The Case of Single Male Cercopithecus pogonias in Troops of Colobus satanas

We report the integration of single male crowned guenons (Cercopithecus pogonias) into troops of black colobus (Colobus satanas). We observed one male Cercopithecus pogonias in three troops of Colobus satanas on 30% of observation days (n = 231). Activities of single males guenons did not differ significantly from those of the colobus with which they associated. Moreover, both species performed simultaneously the same activities more often than expected by chance. Interspecific grooming occurred on several occasions. Furthermore, single male guenons spent as much in time social activities when part of a colobus troop, as they typically do when part of a conspecific group. Unlike solitary male crowned guenons, which are silent, a male that is integrated into a troop of colobus is vocal and emits social alarm calls to which colobus monkeys respond. During the single file movements of colobus troops, single male crowned guenons were integrated in the core of the troop and used the same branches at the same height with the colobus. Thus, the life of a single male crowned guenon with black colobus was social. We suggest that the main benefits that he gained is the possibility to live in a social context. Social interactions could be the key element to explain why single males Cercopithecus pogonias join troops of monkeys so different from their natal groups.     

Adult male replacement and social change in two troops of hanuman langurs (Presbytis entellus) at Jodhpur,India

I studied the process of adult male replacement and social change in two one- male troops (B20 and B21) of hanuman langurs (Presbytis entellus)at Jodhpur, India. Male-male competition lasted for about 6 months before the successful takeover of one troop (B20). During that period, five adult males from three neighboring bands (AMB7, AMB9, and AMB10) and a resident male of a neighboring troop (B21) were involved in taking over the troop. The latter male also copulated with six females during his interim residency, which suggests that he may have opportunistically maximized his mating chances with females of a neighboring group. During an intertroop interaction, a 14-month-old female infant of the other troop (B21) was fatally attacked by an adult female of the first troop and the infant eventually died. The attacker may have taken advantage of the disorganization created by male-male competition, perhaps to eliminate a future food competitor. In addition, the first troop gained an additional feeding area from the other troop’s range; it included a sleeping site and a waterhole, indicating that territorial fights during social instability may have led to the expansion of the winner’s resource area.     

On the dynamics of the hanuman langur populations of jodhpur (Rajasthan,India)

The city of Jodhpur (26°18′N, 73°8′E) supports a population of about 900 hanuman langurs (Presbytis entellus) divided into 24 bisexual troops and 12 all-male bands in an area of 60 km². This population has been censused from 1968 to 1978. Over this period the population of bisexual troops has remained stable around 700, while the population of all-male bands has increased from 160 to 230 individuals. The bisexual troops show a tendency towards a reduction in population growth rate with increasing troop size, with troops over 50–60 tending to split. Very small troops may grow by large scale immigration. Although a number of male changes and mortality through infanticide have been recorded, there is no evidence of a regular periodicity in the occurrence of initial, growth and mature phases in the life history of a bisexual troop. Unlike the bisexual troops, the all-male bands show no tendency towards a reduction in growth rate with the increase in band size, but show a continuous growth of band size over the study period. Langurs of Jodhpur rely heavily on cultivated fields for their sustenance. This cultivation has been on increase over the study period, and since the males invade cultivation more readily, they may have been able to take fuller advantage of these increasing resources and affect a population increase that has not been possible for the bisexual troops.