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.     

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.     

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.     

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.     

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.     

Analysis of population changes by measurement of body weight in the Koshima troop of Japanese monkeys

Measurements of the body weight of monkeys in the Japanese monkey troop on Koshima islet in southern Japan have been made since 1970. Population changes in the troop have been recorded since 1952. The population changes were further analyzed on the basis of the body weight changes of the troop members. The recent history of the Koshima troop can be divided into three periods differing according to conditions of artificial feeding: (1) a semi-wild period (SW Period, 1952–63); (2) an artificial feeding period (AF Period, 1964–71); (3) and a restricted artificial feeding period (Non-AF Period, 1972–77). The AF Period represented a period of population growth, whereas the Non-AF Period was a declining one when the population density of the troop was roughly ten times that of most wild troops. These population changes, i.e., changes in population parameters especially in the population declining phase, could be fairly well understood from the general features of the body weight changes of the troop members, the slow body weight growth, elevation of age at first birth, and small size of adult females. Reproduction and survival were affected by seasonal and yearly changes in body weight. The pattern of changes in body weight displayed seasonal peaks in the autumn and spring. Of these, the former suggests that fruit eating in the autumn is an important factor in the reproduction and growth of the monkeys.     

Some aspects of infanticide and intermale competition among langurs,Presbytis entellus,at Dharwar,India

Conspecific infanticides by male langurs depresses population growth in their densely populated habitats. An infanticidal model developed from demographic parameters obtained in the Dharwar study area predicts an almost stable population. On the other hand, a non-infanticidal model predicts a population growth by 2.6% per year. The effect of frequent troop usurpation and infanticide on the control of population growth must be strong if the natality rate is high. For subadult and juvenile surplus males, it must be difficult to survive and to mature in the all-male party in its poor habitat. However, from calculations for living adult males and the number of troops at Dharwar, most adult males are thought to be able to obtain a troop within five years of first challenge of usurpation. Dominant males do not always take over a troop containing more females than do subordinate males. These simple assessments require further intensive field studies to determine the precise differences between infanticidal and non-infanticidal populations and whether or not dominant males make a greater genetic contribution than subordinate males to subsequent generations.     

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.     

Birth-season variation in Japanese macaques,<Emphasis Type="Italic"> Macaca fuscata</Emphasis>

Japanese macaques, Macaca fuscata, exhibit an annual reproductive cycle that apparently is maintained intrinsically. Translocation of nine troops to new latitudes within the northern hemisphere has had minimal effect on the timing of birth seasonality in these troops; translocation of one troop to the southern hemisphere has resulted in a 6-month forward displacement of birth seasonality in this troop. Limited available evidence indicates that, in the latitudinal zone between Toimisaki (31°22′N) and Kinkazan (38°17′N), mean birth date in in-situ troops becomes earlier as latitude of troop localities increases; the same relationship between mean birth date and latitude apparently does not apply to in-situ troops south and north of the Toimisaki–Kinkazan latitudinal zone. Within the Toimisaki–Kinkazan latitudinal zone, earlier mean birth dates at higher latitudes may permit infants to achieve an adequate level of development before the earlier onset of poor winter food conditions. South of the Toimisaki–Kinkazan latitudinal zone, winters are relatively mild and may be less of a factor in infant survival; north of this zone, poor winter food conditions persist so long that earlier infant births may be maladaptive. Electronic Publication     

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.     

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.     

Population and social dynamics changes in ring-tailed lemur troops at Berenty,Madagascar between 1989 - 1999

In the present study, we recorded all births, immigrations, deaths, and emigrations for a population of ring-tailed lemurs at Berenty Reserve, Madagascar, between September 1989 and August 1999. In September 1989, three troops (C, B, and T) inhabited the study area of 14.2 ha. During the 10-year period, eight troop divisions, six evictions of females, and three troop takeovers of ranges by other troops occurred in and around the study area. Consequently, in August 1999, the number of troops in the same area increased to six (CX, C1, C2A, C2B, T1, and T2). The number of lemurs aged >1 year increased from 63 to 82, which resulted from 204 births, 58 immigrations, 125 deaths, and 118 emigrations. Of the 204 newborn lemurs during the study period, 103 died, 44 emigrated outside the study area, and 57 remained within the study area. The total number of lemurs that emigrated from natal troops was 69 (54 males and 15 females). Natal males left their troops around the age of 3. Non-natal males changed troops after a tenure varying from 1 to 7 years. Survival curves showed a fall in survival rates of both sexes to < 0.5 between the ages of 2 and 3. For females, the survival rate gradually decreased to < 0.2 at the age of 9. On the other hand, due to emigration, the survival rate of males could not be determined after the age of 5 yr. Since some males attained high-rank at the age of 6 – 10 yr, the prime age for male ring-tailed lemurs is thought to be around 7 – 10 yr. Ring-tailed lemurs are essentially female philopatric, because all cases of females leaving natal troops resulted from troop divisions or forced evictions. Such social changes may have resulted from competition among females. All cases of troop divisions or evictions occurred in larger troops consisting of ≥20 lemurs, and only a few females could rejoin their troops. When males joined such a female-group, a new troop was formed. Although promoted by an increase in population, frequent emigrations of females from original troops are the characteristics of ring-tailed lemurs at Berenty.     

Survey of the black howler monkey,<Emphasis Type="Italic">Alouatta pigra</Emphasis>, population at the Mayan site of Palenque,Chiapas, Mexico

A survey of the population of the black howler monkey (Alouatta pigra) present at the Mayan site of Palenque was conducted during 2000. A total of 911 man/hours, spread over 112 days were spent surveying the 600 ha area of pristine forest at the site for howler troops. We detected the presence of 136 individuals of which 131 were members of 20 troops, the rest were 3 solitary adult males and 2 adult males travelling as a pair. Ecological density was estimated at 23 individuals/km². Mean troop size was 7.0 individuals and it ranged from 2–12 individuals; 60% of the troops were multimale. All sighting of howler monkeys were in evergreen rain forest and 75% were in trees ≥20 m in height. The reported densities and mean troop size are higher than those reported for the species in Guatemala and in central Quintana Roo, Mexico. The vegetation of the forest contains tree species reported to be used by species ofAlouatta in the Moraceae, Sapotaceae, Leguminosae, and Lauraceae plant families. Protection of a large perimeter area (ca 1700 ha) around the archeological site by the Mexican government ensures the conservation of the forest and of the black howler monkey population present at the site.     

Population structure,demography, and dynamics of mountain baboons: An interim report

Counts of 61 baboon troops (Papio cynocephalus ursinus) at four localities in the Drakensberg mountains confirmed earlier reports of a small mean troop size. This troop size of 22.49 animals changed neither with latitude nor elevation. Data from two of the sites suggested that population density increases from south to north, while a working assumption of 2.5 animals/ km² allowed us to set the population size at 7,540 animals, living in 335 troops. Both the adult sex ratio of 2.07 females/male and the immature/ adult female ratio of 1.17 were unaffected by troop size. Repeated counts from nine known troops revealed that the population is at equilibrium. © 1995 Wiley-Liss, Inc.     

Patterns of movement by baboons in the Drakensberg mountains: Primary responses to the environment

The movement patterns of two chacma baboon troops in the Drakensberg mountains of South Africa were studied over an 18-month period. One of these troops (HIGH) lived on the escarpment slopes (1920–2980 m), while the other (LOW) lived primarily in the montane belt (1560–2000 m). In this paper we analyze the effects of environmental variables on day journeys and home range use. We found the troops to be similar in the temporal patterning of their daily movement, in the resistance of travel speed to variations in local climatic conditions, and in their response to the nutritional bottleneck at the end of the austral winter. While both troops traveled farther as food became scarcer, HIGH troop undertook longer day journeys than LOW troop. This was probably due to the interactive effects of absolutely lower food availability and lower ambient temperatures. HIGH troop traveled farther when food was scarcer and when minimum temperatures were higher. Neither variable accounted for variance in the day journeys of LOW troop. Both troops used some areas of their home ranges preferentially. For LOW troop this preference followed the spatial distribution of food. While the same was generally true for HIGH troop, its home range use was complicated by the adverse temperatures of winter and spring. It abandoned the higher, more productive part of its range once temperatures became severe, returning only after minimum ground temperatures had risen above 0°C. These usage patterns differ from those described for eastern African baboon populations and reflect both the severity and sesonality of climate and the poor quality and extreme dispersion of food that characterize these southern mountains.     

The effects of food sources on Japanese monkey home range size and location, and population dynamics

The effects of supplemental feeding by tourists on wild Japanese monkey's home range size and location, and troop size and composition were studied for two monkey troops, Troop A and Troop B, living along the Irohazaka loop road, Nikko National Park, central Japan. Changes were documented based on data gathered from 1982 to 1996 by the use of radio telemetry. Troop A's home range size shrank and changed from separate winter and summer ranges to a single, year-round home range, with its core located in a high elevation area where supplemental feeding by tourists was heavy. Troop B's home range also shrank and shifted to a lower elevation where supplemental feeding by tourists was heavy. Troop A's population size increased between the winters of 1983–1984 and 1990–1991 in conjunction with an increase in human encounter rates, and then decreased. Troop B's size increased until the winter of 1993–1994, and then decreased. The instability of troop size between 1993 and 1996 may be explained by documented factors such as a decrease in the adult sex ratio, an increase in the infant-female ratio, and an increase in juvenile mortality and/or emigration, all of which may have been influenced by supplemental feeding by tourists.     

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 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.”     

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.     

Social organization of black-and-white snub-nosed monkeys (Rhinopithecus bieti) at Deqin, China

Data on social organization of two bands of black-and-white snub-nosed monkeys (Rhinopithecus bieti) were collected when the monkeys were crossing an open spot at Nanren and Bamei (northwest of Yunnan, China) using a sampling rule where individuals within one social unit are spatially closer to each other than individuals between social units. The typical pattern of social organization in this sample was multiple adult females (AFs) and their offspring with one adult male (AM) in a one-male unit (OMU), similar to that of many other colobines. In such units, on average one male is associated with 4.0 AFs and 2.5 of their offspring. Moreover, there are multimale/multifemale units and monogamous units besides OMUs. All bisexual units traveled together with at least one all-male unit as a cohesive band. In two bands of monkeys, 87% of AMs in bisexual units were within OMUs, 7.8% within monogamous units and 5.2% within multimale, multifemale units. In the Bamei band, 6.7% of AMs were in the all-male unit. The size of OMUs in the Nanren band was larger than that of the Bamei band, with more AFs and juveniles, which may be related to better conservation in the Nanren band's habitat. For the Nanren band, the average number of AFs in OMUs varied across time, increasing from 4.3 in 1994 to 5.1 in 2001, and then decreasing to 3.8 in 2005. This article suggests three possible explanations for this variation, but more data are needed for these hypotheses to be tested.