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.     

Association and social interactions between strangers and residents in bonobos (Pan paniscus)

This study reports on close spatial association and repeated behavioural interactions between two strange adult male bonobos with residents of another community. Over a period of 12 months one of the two males developed friendly social relations to some of the females and other residents, which were indistinguishable from those existing between co-residents. Aggression by resident males against the strangers decreased but the former remained intolerant. The strange males appeared at a time when the number of adult resident males was lower as in the years before and when the adult sex ratio (number of adult females per male) was higher as in the years before. Using definitions from studies on dispersal patterns of male gorillas (Harcourt, 1978) and female bonobos (Furuichi, 1989) the spatial association between the two strange males and residents could be described as male transfer.     

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.     

The demography of Howler Monkeys (Alouatta palliata) on Barro Colorado Island,Panamá

Chronological ages of Alouattaon Barro Colorado Island (BCI) were estimated from longitudinal dental wear. Combining these data with visual censuses, we approximated the study population’s age profile. A stable model was then constructed; from it we derived agespecific mortality rates. Mortality of immature animals is high, with 88% of the males and 65% of the females dying before 5 years. Adult mortality is low until 11 years, when it accelerates. The average adult life span is 16.6 years for males and 15.5 for females. The maximum life span is over 20 years. A pubertal male growth spurt occurs from 3 to 5 years, at which time females are primiparous. Sexual size differences develop primarily during this time. The 1976 age profile had anomalously few animals aged 7 years and males aged 8–9 and 15–16 years. With corroborating evidence, we hypothesize that these deficits resulted from excessive rainfall in 1963 and 1971, which reduced the fall fruit crop and led to a high juvenile mortality. Within half of our study troops, some adult males have nearly identical ages, suggesting an active process of agemate coalition. We hypothesize a form of kin selection, wherein peripheral male cohorts from the same natal troop have greater survival and social success than solitary animals. The stable model suggests an annual growth rate of 1.5% during the 1970s. Estimates of 16.7 and 4% for the previous two decades, following a yellow-fever epidemic, imply that the BCI population is becoming stationary. A very different age structure existed on neighboring Orchid Island in 1976, suggesting general food limitation. By contrast, the lowered, but continuing growth on BCI suggests a less drastic limiting mechanism. We hypothesize that intratroop social competition limits population growth by regulating subadult survival rates during food scarcity cycles.     

Who is Responsible for Fission in a Free-ranging Troop of Baboons?

The study troop of chacma baboons (Papio cynocephalus ursinus) at Mkuzi Game Reserve, Zululand, South-Africa, comprised of about 76 members that split into two new troops. The events leading to this troop fission will be described and its possible causes will be discussed. Troop fission among baboons is generally attributed to the withdrawal of low-ranking females from the main group, as a result of the cost of food competition and its effect on their reproductive success. At Mkuzi, no evidence for food competition among females was recorded in terms of rank-related time spent feeding or other time—budget components, feeding-bout length, diet composition or context of female aggression. Moreover, no evidence for rank-related differential reproductive success was found in terms of inter-birth intervals or infant survival. Female mortality was, however, related to dominance rank, with circumstantial evidence suggesting that cause of mortality was predation by leopards. Rate of female disappearances, aggression levels among females, and the percentage of time they spent in proximity to other adult troop members increased after fission. Relatively short inter-birth intervals and extremely low infant mortality rate at Mkuzi resulted in a small number of receptive females at any one time, and therefore in high costs of male sexual competition as expressed in the high levels of male aggression and woundings, both reduced after fission. It is suggested that this troop fission may have been initiated by the resident males, triggered by the high cost of sexual competition, and forced on the females, who were, consequently, subjected to higher risk of predation. The troop fission was preceded by a long process of increasing tendency for sub-trooping. It was initiated by the four resident males who kept a large distance apart from each other, herded oestrous female associates away from others and were followed by other females. The females generally tended to stay close to associates, males and females. These parties were followed by the peripheral and immigrant males who had no female associates, and eventually two distinct daughter troops were formed.     

Group composition,home range,and diet of the maroon leaf monkey (Presbytis rubicunda) at Tanjung Puting Reserve,Central Kalimantan,Indonesia

Maroon leaf monkeys (Presbytis rubicunda) were studied at the Orangutan Research Conservation Project study area at the Tanjung Puting Reserve in Central Indonesian Borneo for approximately 15 months in 1974, 1975, and 1977. Over 250 observation hr involving 300 encounters with the monkeys were logged. Troop size ranged from three to eight individuals in the nine troops surveyed with only one adult male being present in each troop. Home range size varied from 0.33 km² to 0.99 km² and correlated with troop size. Forty-six different food types were observed eaten by the maroon leaf monkeys; of these foods, 52% consisted of fruits, 36% of leaves, and 12% of flowers.     

The variable social organization of hanuman langurs(Presbytis entellus), infanticide,and the monopolization of females

Data from 24 wild populations of hanuman langurs (Presbytis entellus)in south Asia are used to test hypotheses seeking to explain variation in troop structure and the incidence of infanticide. The occurrence of infanticide is associated with a one-male troop structure and not with a high density. The density, predation, and economic-advantage hypotheses, as explanations for the occurrence of one-male and multimale troops, are not supported by the review. However, the monopolization hypothesis is not contradicted; the number of adult males per troop is significantly correlated with troop size and with the number of adult females per troop. Therefore it is suggested that a one-male troop structure will arise if a male is able to monopolize a group of females, a multimale troop if he cannot. One-male troops may predispose to infanticide because of high variance in male mating success and high intermale competition between groups rather than within troops. If female dispersion determines troop structure, it is speculated that females could manipulate males to form a multimale society if the advantages in terms of infant survival and intertroop conflict exceeded the costs in terms of not producing infanticidal “sexy sons.”     

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.     

Troop Extinction and Female Fusion in Wild Japanese Macaques in Yakushima

We observed three cases of troop extinction and two cases of female fusion in the wild population of Japanese macaques on Yakushima Island, Japan. Troops P and T decreased in size relatively slowly over a few years until each troop consisted of only three monkeys. Several months later, the remaining adult female of P merged with the adjacent troop S, followed by the remaining female of T. S subsequently also decreased in size and disappeared about 2 years later. In the early stage of troop decline, the mortality rate of adult females was as low as in a growing troop, but the birthrate was quite low. In the late stage of troop decline, the mortality rate increased and the birthrate remained low. An important factor leading to troop extinction may be an increase in population density and the resulting increase in intergroup competition. During the period when P and T declined and ceased to exist, the range of the adopted troop shifted to cover their previous ranges. In the fused troop, there was no severe aggression directed towards the immigrant females or harassment from residents of the adopted troop and there was affiliative social interaction between the immigrant females and resident members. These results agree with previous reports on female fusion: it occurs when the shrinking group consists of one or no adult member, and the immigrant females are not at a severe disadvantage in their adoptive group. A possible benefit for immigrant females is to avoid disadvantage of one-adult group in conflict with conspecifics. A possible cost for immigrant females is transfer to the other troop or to unfamiliar area or both. The cost to transfer to another group may not be high because the members of the adoptive troop are relatively tolerant to immigrants. The cost to transfer to unfamiliar range may be minimized by immigration to the troop whose range shifted to the immigrants' former range.     

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.     

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.     

Does troop size of wild Japanese macaques influence birth rate and infant mortality in the absence of predators?

For the wild Japanese macaques of Yakushima and Kinkazan Islands, we analyzed the relationship between the troop size or the number of adult females of each troop, infant/adult female ratio (IFR; crude birth rate), and infant mortality (IM) in habitats with no predators. In Yakushima, IFR was positively correlated to troop size and the number of adult females. In Kinkazan, however, IFR tended to decrease with the number of adult females. This difference may be due to the difference in troop size; i.e. in Yakushima, where troop size was small, IFR may increase with that of troop size, because a relatively larger troop is likely to the advantage in intertroop competition. In Kinkazan, where troop size was large, however, IFR is likely to decrease with troop size, because intratroop competition may increase. Thus, the present data roughly supportWrangham's model of the social structure of female-bonded primates, and suggests that there is an optimal troop size for birth rate (BR). On the other hand, there was no clear correlation between IM and the troop size or number of adult females of each troop.     

Dispersal Patterns Among Olive Colobus in Taï National Park

In Primates, females are more likely to be philopatric than males. However, in some species like Procolobus verus, females or individuals of both sexes disperse. In Taï National Park, Ivory Coast, olive colobus groups are small, with one or two adult males and 6 females. Dispersal is common for juveniles and adults of both sexes. Adult male dispersal is less common than adult female dispersal. Adult females immigrated especially into small, one-male groups indicating that food competition played a role. Furthermore, unknown sexually receptive females visited resident groups and mated with the resident males for a few days before disappearing again. Adult males dispersed when this improved their mating opportunities. All juveniles left their natal groups. The dispersal of juveniles may be a strategy to prevent inbreeding with their parents. Dispersal by juvenile males furthermore seemed to be the result of mate competition. The high dispersal rates, visits by receptive females, and dispersal of all individuals in the population suggest that moving between groups is a strategy that can be used ad hoc in several situations more easily in the olive colobus than in most other primates. The predation risks related to moving between groups were reduced by dispersing in conspecific or allospecific groups and by dispersing to neighboring groups.     

Somatometrical features of Japanese monkeys in the Koshima islet: In viewpoint of somatometry,growth, and sexual maturation

The Japanese monkeys inhabiting Koshima islet were investigated morphologically, and compared with those living in other areas of Japan. The morphological features of the Koshima monkeys are thought to reflect many interrelated factors. Their body size is the smallest inMacaca fuscata fuscata. The lower latitude and the warmer winter of the islet represent the major factors in this small body size. Another physical characteristic is their longer limbs, which may be related to the same environmental factors. The adult males of the Koshima troop show the widest variation, while the variation in the adult females is the narrowest, among the troops ofMacaca fuscata examined. This may be explained either by sexually differentiated growth, that is, males continue to grow considerably after 8 years of age, whereas females do not; or by isolation, that is, males sometimes immigrate into or emigrate from the islet, whereas almost of all the females remain there. It was found that the influence of food limitations on body weight depends on sex and age; babies and juveniles, and adult males did not lose their body weight, but monkeys aged from 3 to 7 years and adult females had lost much weight compared to those of the same ages and sexes measured in 1963 (feeding was not limited). This fact may support the idea of “sex-age-class selection” proposed byMori (1979), but the weight loss pattern could be explained by changes of body composition with growth. The growth of the anterior trunk length, too, was restrained. Sexual maturation in males was delayed by at least one year.     

Life-history characteristics of a wild population of vervets (Cercopithecus aethiops sabaeus) in Barbados,West Indies

Life history data are presented for a population of vervets, Cercopithecusaethiops sabaeus, in Barbados, West Indies. The data were obtained from two habituated troops and from vervets captured during a large-scale trapping program. Individuals of known age from one troop were weighed periodically, and separate growth curves generated for males and females. The mean weight of captured adult females was 3.3 kg; that of adult males, 5.3 kg. The average age at sexual maturity is estimated at 34 months for females and 60 months for males. Vervets give birth throughout the year, but most infants are born between April and July. The average interbirth interval following a surviving infant is 11.8 months. The mortality of juveniles is heaviest between birth and 2 years of age and decreases thereafter. Males emigrate from their natal troops at sexual maturity and one incident of a juvenile female emigrating is reported.     

Social stability and change among forest hanuman langurs (Presbytis entellus)

The social organization of hanuman langurs (Presbytis entellus, Colobinae) was studied in Kanha Tiger Reserve, Central Indian Highlands between 1980 and 1983, followed by six brief return visits over ten years. Censuses of the 7 km² Kanha meadows demonstrated little change in population density and structure between 1982 and 1990; the population was consistently composed of one-male troops and all-male bands. During the return visits the focal C troop extended its known 74.5 ha range by only 5.6%. By 1993 two identified, habituated, adult females remained within the troop. The adult male was resident in C troop for ≥nine years and eight months.     

Sexual Maturation of Female Japanese Macaques Under Poor Nutritional Conditions and Food-Enhanced Perineal Swelling in the Koshima Troop

Provisioned food was severely restricted for 5 years, from 1972 until June 1977, in the insular Koshima troop of Japanese macaques. The monkeys experienced very poor nutritional conditions during this period. The feeding policy was changed in 1977 and food conditions improved, but only in summer. A remarkable consequence of intensive provisioning in summer was the occurrence of perineal swelling in many young females outside of the mating season. Females that matured during the period of poor food conditions showed an extraordinary delay in primiparity and exhibited conspicuous perineal swelling in the summer of 1977. Researchers studied the development of perineal swelling under standard conditions of constant provisioning from 1990 to 1995. First perineal swelling appeared at 5 years of age under standard conditions but appeared at 8 years of age under poor food conditions. The extraordinary delay in primiparity did not occur among females that were 6 years old when food conditions were improved in summer. We deduced that female Japanese macaques have a critical age of 5 to 6 years for sexual maturation. If the critical age is exceeded under poor food conditions, sexual maturation is hard to achieve even with improvements in food conditions. Seasonal changes in the incidence of perineal swelling were related to the period of high body weight. Rapid improvements in food conditions probably caused perineal swelling and activated sex hormones in female Koshima Japanese macaques.     

The differential use of cheek pouches in a troop ofPapio ursinus

Cheek pouches, one of the distinguishing characters of the Cercopithecinae, are structures used for the temporary storage of food. Their size and frequency of use within a given species are related primarily to the amount of conspecific competition for food. In relation to total body size, members of the genusPapio are considered to have relatively small cheek pouches which are said to be used only occasionally to maximize food harvest when local clusters of food are encountered. This investigation represents 165 hours of observation on a troop ofPapio ursinus at Mkuzi Game Reserve in Natal, South Africa. At the time of observation the choice of foods in the home range was restricted and usually found in abundance only in small clusters of trees. Given that conspecific competition would have been pronounced under these conditions, it was thought that any differences in the frequency of cheek pouch use related to age, sex, rank, or the reproductive state of an animal would be readily recognizable. The results indicated that cheek pouches were used differentially throughout the day, with virtually all ages of each sex displaying the same general pattern of maximal cheek pouch use during the middle of the day. In each sex there was an age graded diminution of the frequency of cheek pouch use from juveniles to adults. This trend was more pronounced in males resulting in a substantial sex difference in the use of cheek pouches between adult males and females. In addition, there was some indication that differences in the frequency of cheek pouch use between adult males were correlated with rank. Although a pattern of cheek pouch use and rank was not evident amongst adult females, there was an association between reproductive state and the frequency of cheek pouch use. Overall, body size, dominance, and energetic demands appeared to be the most significant factors underlying the differences in cheek pouch use in this troop.     

Female preferences for male golden snub-nosed monkeys vary with male age and social context

Age is a key factor affecting sexual selection, as many physical and social traits are age-related. Although studies of primate mate choice often consider particular age-related traits, few consider the collective effects of male age. We tested the hypothesis that female golden snub-nosed monkeys Rhinopithecus roxellana prefer prime aged males (10–15 years) over younger and older males. We examined a habituated, provisioned troop during a 3-year study in the Qinling Mountains, China. Prime age males were more likely to be resident males of 1-male units (OMUs) than males of other ages. Since females are free to transfer between OMUs, the number of females per OMU can be indicative of female preferences. We examined the number of females per OMU, and found that it increased with resident male age up to 7–8 years, and declined after 12 years, such that prime age resident males had more females than other resident males. Females also initiated extra-unit copulations with high-ranking prime age males at significantly higher rates than with other males. Nevertheless, females tended to transfer from OMUs with high-ranking, older resident males to those with low-ranking, younger resident males. Thus, females appear to use different strategies when choosing social mates and extra-unit mates (i.e., different social contexts). We speculate that females may perceive early signs of aging in males and trade off the benefits and costs of high rank versus male senescence. This study lays the groundwork for future studies that examine possible direct and indirect benefits of such strategies.     

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.