Twins in free-ranging Hanuman langurs (Presbytis entellus)

Two pairs of twins were observed in free-ranging Hanuman langurs (Presbytis entellus) at Jodhpur, India. Each twin had a preferred nipple for suckling, the preference of which developed during the first 5 weeks of life. The behaviour of the twins was mostly synchronized. In more than 60% of all activity bouts both showed the same behaviour, with “nipple contact” and “playing” prevailing. Differences in physical strength and development were apparent within each pair. While the total amount of allomothering was almost the same, individual weekly scores differed considerably.     

Sociobiology of Hanuman langurs (Presbytis entellus): introduction into the Jodhpur field project

The conceptual development of a longterm project on Hanuman langurs (Presbytis entellus) is described. The joint indo-german project was performed at Jodhpur, India, starting in 1977. Its main aim was to study the interrelations between individual life histories and troop histories. A general overview of the habitat and the langur population is presented.     

Fate of ousted male residents of one-male bisexual troops of Hanuman langurs (Presbytis entellus) at Jodhpur,Rajasthan, India

Little is known about the fate of adult male residents after they are ousted from bisexual one-male troops of Hanuman langurs (Presbytis entellus) in the course of adult male replacements. In a long term study at Jodhpur, Rajasthan, it was possible to reconstruct partial life histories of several ousted residents. One resident was killed during the male change. Ousted residents did not regain residency despite their continued invasions into bisexual troops. It is assumed that the males' chances to take over and to defend a troop are restricted to an age of 9–14 years, when the males are in prime physical condition. One male became solitary for some months while trying to regain residency of his old troop, before joining some “alien” males. As a rule, males are likely to rejoin their own male bands if they are ousted after short periods of residency. If the residency exceeds 3 months then the ongoing structural change in the former band may prevent their reintegration. However, in such cases, ousted residents which belonged to the same band may reunite and mingle with another male band which lacks prime males. Weaned sons may follow their fathers after ousting. In the case of numerous weaned offspring, fathers and sons may together form at least temporary new male bands.     

Male competition and coalitions in langurs (Presbytis entellus) at Jodhpur,Rajasthan, India

During a 15 month study on free ranging langurs (Presbytis entellus) at Jodhpur, Rajasthan, India, 5 adult male replacements were observed as a result of nontroop male invasions into the home ranges of 3 neighbouring one-male troops comprising 16–28 members each. Jodhpur langurs have no breeding season. Periods of instability during resident male changes lasted 11–119 days. Linear dominance hierarchies could be detected within the 3 main rival male bands of 2, 5, and 28–35 members. The respective alphas drove their allies away after their bands succeeded cooperatively at occupying a troop. During gradual replacements interim residencies alternated with multi-male stages. A large band's alpha may have had better chances to win the competition, since adult and nonadult allies functioned as “buffers” in agonistic encounters. The role of kin selection in structuring the composition of male bands and male coalitional behaviour cannot yet be quantified. Tactical “deceit” of powerful males to cause unrealistic expectations and in this way agonistic engagement of less strong males can be ruled out. “Sneaking copulations” is a proximate advantage for subordinate supporters, since they participated in 61.9% of all sexual interactions. Female promiscuity might reflect a strategy to induce male-male competition and thus select for a strong resident.     

Patterns of grandmaternal behaviour in free-ranging Hanuman langurs (Presbytis entellus)

The behaviour of two Hanuman langur grandmothers towards their grandchildren is examined with emphasis on the spatial relationship, the contact behaviour, and vigilance. Interaction patterns with infants and juveniles are analysed separately to disclose the influence of a social partner's reproductive value and the variance of this value on grandmaternal behaviour. The shape of grandmaternal behaviour seems to differ for male and female grandchildren, and it seems to depend on the reproductive value of both, the grandmother and the grandchild. These results are in accordance with sociobiological hypotheses. Evaluation of costs and benefits suggests that grandmaternal behaviour in langurs can be called investment.     

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.     

Father-son recognition in hanuman langurs (Presbytis entellus): A field experiment in India

A juvenile male langur, about 15 months old, was removed from his natal one-male-multi-female troop and reintroduced after 3.3 weeks. In attempts to protect the juvenile, his presumed father, the troop's resident male, charged the primatologist who removed and released the individual. After the second removal, the juvenile was introduced into a neighboring male band. The resident male attacked the band members to keep them away from his females. However, after 3.4 weeks of separation, the father and the son recognized each other, because the resident male showed no aggression towards his son and the latter did not run away from him. Kin-recognition is considered to be a mechanism in structuring coalitions within langur male bands.     

Troop history,female reproductive strategies and timing of male change in Hanuman langurs,Presbytis entellus

Female reproductive data are presented from 9 years of longitudinal observations on two troops of Hanuman langurs (Presbytis entellus) living around Jodhpur, India. On the basis of 89 live births interbirth intervals were calculated to examine the effect of demographic factors on reproductive behaviour and troop composition. Sex of an infant seems to influence the length of intervals which are longer after the birth of female infants at an average of 1.7 months. It is suggested that this may be an outcome of differential maternal investment by allocating more time and energy towards female infants who run a higher mortality risk than male infants, at least up to an age of 27 months. Troopspecific interbirth intervals are influenced by social events. If the last infant is still alive when the next one is conceived, the intervals are significantly longer than after the premature loss of an infant (Bijolai troop: 15.6 vs. 12.1 months; Kailana-1 troop: 16.7 vs. 11.4 months). During undisturbed male tenureship intervals are shorter than after a male change (Bijolai troop: 14.3 vs. 16.0 months; Kailana-I troop: 15.6 vs. 17.5 months). Thus the frequency of male changes can influence the demography of a troop. Furthermore, the data suggest that take-overs are optimally timed by males. New males tend to take over a troop when most of the females are cycling.     

Birth spacing in langur monkeys (Presbytis entellus)

This paper presents 10 years of reproductive data on birth interval length and 5 years of data on reproductive behavior postpartum from a captive colony of gray langur monkeys (Presbytis entellus)housed in Berkeley, California. Birth intervals of females following different pregnancy and nursing schedules are compared. Females whose infants survive to the age of 9 months have a median birth interval of 15.4 months. The experimental separation of mothers from infants for a period of 2 weeks, 6 to 9 months postpartum, had no significant effect on the median birth interval length. Females experiencing a pregnancy failure or the loss of a neonate had median birth intervals of 9.6 and 10.7 months, respectively. These intervals were significantly shorter than the birth intervals of females whose infants survived to 9 months, showing that the presence of a nursing infant delays the female’s time to next conception by approximately 5 to 6 months. Females experienced a median of three estrous periods (two estrous cycles) before conceiving postpartum, regardless of pregnancy outcome or length of infant survival, and females rarely conceived during their first estrous period postpartum. Weaning did not occur until after the mother’s next conception. These data indicate that, in populations of langurs characterized by average birth intervals of 15 to 16 months, the loss of an infant after the age of 5 to 6 months will not accelerate a female’s ability to conceive or shorten the birth interval length. The available data on birth spacing from populations of free-ranging langurs are reviewed. It could not be demonstrated that non-Himalayan populations are characterized by birth intervals which are as long as 20 to 24 months. Rather, it is suggested that female langurs inhabiting seasonally arid sites, such as Jodhpur, Abu, and Dharwar, may be capable of producing infants on the average of every 15 to 16 months. Flexibility in the timing of births and the lack of well-defined birth seasons at these sites may be explained by this species’ dietary and digestive adaptations. Additionally, data on birth spacing and the age of missing infants from the above field sites, where it has been suggested that infanticide following changes in male leadership occurs habitually, do not lend support to the sexual selection hypothesis of infanticide as proposed by S. Hrdy (1974, 1977).     

Grooming site preferences in female langurs (Presbytis entellus)

Factors influencing grooming site preferences in adult female Hanuman langurs (Presbytis entellus) were investigated. The females belonged to a free-ranging harem troop (Jodhpur, India) and were observed for 569 hr by focal-female sampling. Decisive factors for grooming site preferences were the following: autogrooming was determined mostly by site accessiblity. Allogrooming was significantly concentrated on parts that are inaccessible to the groomee. Close female kin groomed significantly longer, more frequently, and more precisely at inaccessible body parts. Lower-ranking females were groomed significantly less often and more briefly but also more precisely at inaccessible parts. However, the latter might be due to a lower-ranking subjects desire to face away from the higher-ranking groomer in order to avoid eye contact. The data suggest that the groomee determines the sites being groomed.     

Weaving a tight social net: Allogrooming in free-ranging female langurs (Presbytis entellus)

We studied grooming among adults of a one-male multifemale troop of free-ranging Hanuman langurs (Presbytis entellus)living near Jodhpur, India, for 9 years. The 11–13 females devoted about 6% of their day to allogrooming. Adult males, whose tenures averaged 2.2 years, were transient figures in the troop's history, as reflected by their rather peripheral role in the grooming network. Females groomed males 4–40 times more frequently (1006 episodes) than vice versa- (176 episodes). Adult females received 97% of all grooming from other adult females (6655 episodes). Although females exhibited an age- inversed dominance hierarchy, they did not compete for grooming access to particular troop mates. Dyads of all possible rank differences occurred as frequently as expected: 51% of grooming was directed up the hierarchy and 49% down it. Young, high- ranking individuals gave and received significantly more grooming than the oldest, low- ranking females did. The pattern seemed to be influenced by kin selection because of the presumably high degree of female relatedness. They invested most in troopmates with the highest reproductive value, i.e., the youngest individuals. This trend was coupled with a preference of closest kin (mothers and daughters). Reciprocity was the outstanding feature since all adult females groomed and were groomed by all others. Such a tight social net might establish the necessary cohesion during frequent territorial disputes with neighboring troops.     

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.     

The social organization of forest hanuman langurs(Presbytis entellus)

The social organization of hanuman langurs (Presbytis entellus;Colobinae) was studied in Kanha Tiger Reserve, Central Indian Highlands, for 2300 hr (1980–1985), in a mosaic of moist deciduous forest and anthropogenic meadow. The langur population density was 46.15/km ² and the mean troop and band sizes were 21.7 and 14.0, respectively. Of 14 troops, 13 were one-male and 1 was trimale. The population adult sex ratio was 1:2.5. The majority of female sexual solicitations was directed toward the harem male. The birth season was December to May, with an estimated gestation of 171–224 days. A review of langur reproductive seasonality suggests that breeding throughout the year is confined to those populations able to exploit human food sources. Mortality during the first year of life was 40%, including infanticide. A significant positive correlation was found between the age of an infant at death or disappearance and the mother’s subsequent interbirth interval. Five cases of social change are described, including female transfer, one-male to multimale change, troop formation, and gradual and rapid replacement of troop males. Takeover-associated infant killing by band males, in an undisturbed moderate-density population, supported the sexual-selection/infanticide hypothesis but not the social-pathology hypothesis. However, it could not be directly confirmed that an invading infanticidal male gains a reproductive advantage. The male tenure of harems was estimated to be 45 months.     

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.     

Infanticide and juvenilicide in Hanuman langurs (Presbytis entellus) around Jodhpur,India

Ten cases of infant killings and 2 cases of juvenile killings were observed in two troops of Hanuman langurs, (Presbytis entellus) around Jodhpur, India. Fatal attacks on infants and juveniles are classified into four categories. The process of infanticide was observed from start to end and is described in detail for 3 cases. The age of the victims ranged from 0.2 to 48 months. The interbirth interval among females whose infants were killed is significantly shorter compared to females whose infants survived. In ourt study, 7 cases support the reproductive advantage hypothesis, that infanticide is an adaptive behaviour to increase male reproductive success. The remaining 5 cases do not fit into the reproductive advantage hypothesis. In these cases, victims are over 8 months old, and as such their deaths could not shorten the interbirth interval. It appears that by killing older infants and juveniles the males obtain an advantage in resource competition for their offspring. An alternative is that new males chase or peripheralise the older infants and juveniles, which leads to 97% predominant uni-male troop structure in Jodhpur.     

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.     

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.     

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.”     

Sex Differential Behavior of Immature Hanuman Langurs (Presbytis entellus) in Ramnagar, South Nepal

We investigated sex differences in the social behavior of immature Hanuman langurs (Presbytis entellus) in the light of sex-specifically different life-courses and Hanuman langur characteristics, such as the individualistic dominance hierarchy and the rarity of intragroup coalitions among adult females. We observed four immature female and four immature male langurs—all members of the same free-ranging multimale multifemale group in Ramnagar, South Nepal—from November 1992 to February 1993 for 288 hr via focal-animal and instantaneous sampling techniques. Immature females spent significantly more time in proximity to other group members than immature males did. They had more physical contact and groomed more. Other immature females were their preferred social partners. Immature males also preferred like-aged females. They restricted their relationships with other immature males to proximity and occasional grooming. Monitoring was directed especially toward adult males. Female behavior can be interpreted as oriented toward integration into the female social network and their age-inverted dominance hierarchy. Males seem to prepare for leaving their natal group and for future strong intrasexual competition.     

Take-over and infanticide in South Indian Hanuman langurs (Presbytis entellus)

A take-over and infanticide were observed in a South Indian population of Hanuman langurs (Presbytis entellus). A one-male, bisexual troop of langurs (Troop B1) was attacked by a band of two adult males and one subadult male. During the take-over, the resident male and one elderly female were driven from the troop. The youngest infant, about two months old, was fatally wounded by an attack from the dominant incoming male (S1). However, other young infants in Troop B1 were left unharmed. All three incoming males remained in Troop B1 but the troop appeared to remain functionally one-male, as S1 did not allow the other adult male to copulate. There was no sign that S1 attempted to eject the other two males from troop B1. © 1993 Wiley-Liss, Inc.