Troop Males Research Articles

Association patterns and affiliative relationships outside a troop in wild male Japanese macaques, Macaca fuscata, during the non-mating season

In many mammals, males disperse more than females. Although males in some male-dispersing species form all-male groups, little is known about the association patterns or social relationships among them. Studies on male–male social relationships have primarily focused on competition over fertile females, but affiliative relationships among males have attracted much interest recently. I suggested a novel method for the classification of males based on their behaviour by principal component analysis (PCA), and investigated association patterns, and affiliative relationships in male Japanese macaques, Macaca fuscata, during the non-mating season. I observed 12 wild male Japanese macaques for 809 h during the non-mating season. The number of other animals within visual range, the amount of time that males spent in the vicinity of other animals, and the frequency of grooming interactions were examined. I classified males into two distinctive clusters (Cluster 1 and 2) according to their association and interaction patterns. Cluster 1 males associated with females and participated in grooming with them. Cluster 2 males had less visual encounters with females and did not groom them. Cluster 2 males showed proximity to other Cluster 2 males in all possible dyads. Although Cluster 2 males showed less proximity to each other than Cluster 1 males did, they frequently exchanged grooming among themselves. Cluster 2 males groomed Cluster 1 males more frequently than they were groomed by them. These results suggested that Cluster 1 were troop males, and Cluster 2 were non-troop males. Cluster 2 males had less opportunity for social interaction than Cluster 1 (troop) males, and they may form all-male groups. Males in all-male groups engaged in more frequent grooming than troop males. In addition, they groomed troop males more frequently than they were groomed. These results suggest that males could be separated by their behaviour. Male–male affiliative relationships might be influenced by within-group potential competition and imbalanced grooming appears to cause troop males to tolerate non-troop males which might be immigrated in near future.

Random Sample

The Disney film Chimpanzee tells a scientifically accurate story about an orphan chimp that was adopted by one of his troop's males—in fact, "the main plot was decided by the chimps," says primatologist Christophe Boesch of the Max Planck Institute for Evolutionary Anthropology. A noted Maya epigrapher sees the cottage industry in Maya 2012 predictions as a "once-in-a-lifetime opportunity" to inform people about the true role time played in the Maya's lives. And this week's numbers quantify the percentage of pediatric drug trials and the number of Chinese family names.

Troop Takeover and Reproductive Success of Wild Male Japanese Macaques on Yakushima Island (Macaca fuscata yakui)

Troop takeover is common in one-male primate groups, but there are few reports in multimale groups. Japanese macaques (Macaca fuscata) form multimale groups and males commonly join troops at the bottom rank. On Yakushima island, however, where group size is relatively small, entrance into groups at the alpha position is also observed. This paper reports on the general features of troop takeover, on the predictors of takeover events, and on the reproductive success of takeover males. Troop takeovers occurred only in the mating season; nontroop males (NTMs) did not cooperate with each other; former alpha males were rarely expelled from the troop; new alpha males did not commit infanticide; new alpha male tenure in the group was usually less than two years. Logistic regression analysis showed that the number of NTMs associating with a troop predicted the occurrence of troop takeover. Paternity discrimination revealed that 33.3% (3/9) of takeover males succeeded in siring offspring. Contrary to this low success rate, binary logistic regression analysis revealed that the takeover males can expect higher reproductive success compared to troop males. Entering a troop and out-competing the alpha male is one of many available strategies to attain reproductive success in male Japanese macaques.

Case study of a one‐sided attack by multiple troop members on a nontroop adolescent male and the death of Japanese macaques (Macaca fuscata)

An adolescent wild male Japanese macaque (Macaca fuscata), following Kinkazan A troop, was attacked one-sidedly by multiple members of the troop. The victim was identified as PI, and was estimated to be seven+/-one year old. The aggressive interaction was recorded by video camera until the end. Although at least 16 troop members approached PI more than once, only three males (one adult, two adolescents) of A troop attacked him. PI kept crouching throughout the attack, then escaped to the shore and dived into the sea. The interaction continued for more than one hour. PI was found dead a few hours after the end of interaction. The damage caused by the assailants was not the direct cause of PI's death; it was due to hypothermia caused by drifting in the sea. PI's life history was reconstructed from past records. PI was a normal adolescent male who migrated from an all-male group around B1 troop and started ranging around A troop. The aggressive interaction is believed to be a typical example of conflict between troop males and a nontroop male. The interaction period was very long compared with previous reports on such conflicts among Japanese macaques. PI kept crouching in open areas, exposing himself as a potential competitor for the resources of the troop, and did not show any submissive or reconciliatory behavior toward the troop males. This may be why the troop males did not stop the attack.

Female Defensibility in a Small Troops of Japanese Macaques vis-à-vis Nontroop Males and Copulation on the Periphery of the Troop

I provide data compiled over 4 yr on the mating behavior in small troops of wild Japanese macaques on Yakushima Island. The key parameters are the number of sexually receptive females, the number of nontroop males (NTMs), and copulation on the periphery of the troop. I analyzed the following aspects: 1) changes in the proportion of copulation with high-ranking males (HRMs) and NTMs, 2) variations in factors such as fluctuation in the number of sexually receptive females and troop males and their effects on the number of visiting NTMs, 3) the effect of attempted interruption of mounting series by other males, and 4) some aspects of copulation on the periphery of the troop. Throughout the study, 56% of the total number of females mated most frequently with the α-male in a single mating season. However, the relative mating success of HRMs varied over the years and between individuals. The number of visiting NTMs varied depending on the number of receptive females and troop males. Females tended to mate with the NTMs when they appeared around their troops. The direct effect of interruption of the mounting series by other males is equivocal. The females mated with the low-ranking males (LRMs) and NTMs on the periphery of the troop, which increased the possibility of mounting series ending with ejaculation. Females actively sought opportunities for copulation on the periphery of the troop by moving there or initiating close proximity with LRMs and NTMs there. On Yakushima Island, the mating success of HRMs was not always as high as that predicted by the priority of access model. The injury status of the HRM, the number of visiting NTMs, and female choice are all considered to influence a male’s mating success.

Social relationships of male Japanese macaques (Macaca fuscata) in different habitats: a comparison between Yakushima island and Shimokita peninsula populations

Wild troops of the Japanese macaque (Macaca fuscata) live in various habitats within the Japanese archipelago. The social relationships of males can be elucidated through intraspecific comparisons of social conditions between populations of different habitats. Previous research has shown that males associate with one another more frequently on Yakushima island than on Kinkazan island, whereas male–female and female–female relationships did not differ between populations that occupy the two different habitats. However, this study did not compare the social relationships of males between habitats during the mating season, when males are expected to compete for fertile females more aggressively within and between troops. The present study compared male behavior in Yakushima island and the Shimokita peninsula populations during the mating season and found that males exchanged affiliative behaviors with one another more frequently on Yakushima island. The socionomic sex ratio (SSR: adult males/adult females) was higher in the Yakushima island troop, and this troop was involved in inter-troop encounters more frequently. In Yakushima island, more non-troop males were observed around the troop and were frequently involved in agonistic interactions with troop males. Yakushima island troop males are likely to affiliate with one another to reduce tension among themselves, in order to cooperatively attack neighboring troops and non-troop males that appear to copulate with fertile females.

Affiliative relations among male Japanese macaques (Macaca fuscata yakui) within and outside a troop on Yakushima Island

Male Japanese macaques (Macaca fuscata yakui) in a troop on Yakushima Island frequently groom other males. However, previous studies have not compared the social relations of troop males to those of non-troop males. I followed all troop males and non-troop males in and near a troop during a mating season and during the following non-mating season and recorded their neighbors, grooming, and agonistic interactions. Comparisons of the social relations of troop males and non-troop males with other troop members revealed that grooming and agonistic interactions with females during the mating season were similar between troop and non-troop males. However, troop males groomed each other more often and had fewer agonistic interactions among themselves than did non-troop males. Compared to what occurred in the mating season, troop males groomed females less often and exchanged grooming bouts more often with other troop males during the non-mating season. One non-troop male groomed females more frequently than did any troop male in both seasons, and this male groomed troop males more frequently than did any troop male in the non-mating season. This male immigrated into the troop during the following mating season. Regardless of their competition with respect to reproduction, male Japanese macaques on Yakushima Island maintain affiliative relations, probably to cooperatively defend fertile females from non-troop males.

Male Masturbation in Free-Ranging Japanese Macaques

We studied masturbatory behavior of males over a 14-mo period in free-ranging Japanese macaques (Macaca fuscata yakui) on Yakushima island, Japan. We distinguished 2 types: (1) masturbation only and (2) masturbation with final ejaculation. Both types occurred in all 15 adult troop males. We calculated masturbation only as an index (M/h). It occurred over the whole year; however, it was more frequent during the mating than during the non-mating periods. Masturbation with ejaculation was restricted to the mating period. Male mating success correlated positively with social status. Both rate of masturbation only and masturbatory ejaculations increased as male social status and male mating success declined. Lower-ranking males had higher rates of masturbation only, i.e., they spent more time masturbating than higher-ranking males did. Lower-ranking males also had higher proportions of ejaculates for masturbation, while higher-ranking males used most of their ejaculates for mating.

The interaction of male and female reproductive strategies and paternity in wild Japanese macaques, Macaca fuscata

Japanese macaques reside in large, mixed-sex social groups in which various reproductive strategies of both sexes operate simultaneously. This report represents the first study combining behavioural and genetic data to examine the interaction of male and female reproductive strategies in primates (N=15 adult males, N=15 adult females, Yakushima Island, Japan). During one mating season, socially dominant males monopolized most female matings. Furthermore, the six offspring sired by troop males were more likely sired by higher-ranking males than lower-ranking males. Nontroop males sired three additional offspring in the troop. Lower-ranking troop males avoided direct competition with higher-ranking males by engaging in sneak copulations with females outside of the presence of other males. Also, females expressed mate choice behaviour towards multiple males of various dominance ranks. Thus, the female strategy of attempting to mate with multiple males conflicted with the mate-guarding strategy of high-ranking males. Despite some female mate choice for mid- and low-ranking males and alternative male mating tactics by subordinate males, high-ranking males were able to monopolize most, but not all, within-troop mating and paternity. This result was due in part to the low number of females mating at the same time. The mean number of females displaying mating behaviour per day was 2.42 (range 1–5), and higher-ranking males more successfully monopolized females on days when fewer females were mating. The number of females mating simultaneously influences the outcome of reproductive conflicts between the sexes.

Influence of fluctuation in the operational sex ratio to mating of troop and non-troop male Japanese macaques for four years on Kinkazan Island, Japan

Taking advantage of a marked yearly fluctuation in the number of estrous females, I studied the differences in mating success between troop males and non-troop males in an unprovisioned group of Japanese macaques. Fluctuation in the defendability of estrous females by troop males, as predicted by the operational sex ratio (the number of estrous females per troop male), strongly affected the mating with ejaculation (successful mating) per observation day of both troop and non-troop males. When operational sex ratio was low, troop males monopolized successful mating inside the troop. No successful mating of non-troop males was observed inside the troop. In contrast, both troop and non-troop males were able to mate often inside the troop when operational sex ratio was high. These findings suggest that troop males obtained the benefit of secured successful mating in the troop because troop males could mate successfully even in mating seasons with a low operational sex ratio, and the chance of successful mating for non-troop males will increase as the ability of troop males to monopolize estrous females decreases.

Contrast of Estrus in Accordance with Social Contexts Between Two Troops of Wild Japanese Macaques on Yakushima.

I observed estrus in two adjacent troops (A-troop and M-troop) of wild Yakushima macaques during the 1984 mating season. A-troop females showed short and regularly cycling estrus periods, seldom came into estrus simultaneously, and scarcely failed to mate when they were in estrus. In contrast, females of M-troop, which went through troop takeovers during the mating season, demonstrated mate competition and subsequent estrus prolongation. Early in the mating season, constantly four to seven females of M-troop were in estrus simultaneously, competed for troop males (TMs), and some of them failed to mate. Subsequently many non-troop males (NTMs) approached to the troop, estrous females mated with them frequently, and mating harassments by females decreased. Among those NTMs, one dominated all TMs, which was referred to as a troop takeover. Successively three other NTMs dominated both all TMs and the former dominant NTM, and in total, four troop takeovers occurred in M-troop during this mating season. The females continued to be in estrus during the whole episodes. All the conceived females also showed prolonged postconception estrus, and they were neither inactive nor less attractive to males. It was suggested that M-troop females increased their opportunity to mate with NTMs by estrus prolongation, at the cost of female-female mate competition, to incite active male intertroop movement and subsequent troop takeovers.

Infant killing, wounding and predation in Eulemur and Lemur

Infant killing by primates is highly controversial. Sexual selection of infanticidal males has been disputed, especially for seasonally breeding species, in which death of an infant does not advance conception of the next infant. We report attacks, infants found wounded, and predation in seasonally breeding Eulemur and Lemur at Berenty, Beza Mahafaly and Duke University Primate Center, and review cases seen elsewhere. Observed attacks leading to wounds or death conservatively total twelve by extratroop males, two by troop males, and seven by troop females. Eulemur are occasional vertebrate predators, whose prey includes infant Lemur catta. Wounds inflicted by lemurs are usually abdominal canine slashes or bites to the head, with rare eating, a pattern distinct from carnivore and raptor kills. Infant killing as inferred from corpses is more frequent than previously thought, but still rare. Adaptive advantages of killing plausibly include eliminating resource competitors of females, and sexual selection on males.

The behavior of peripheral males during the mating season inMacaca fuscata

The mating season behavior of peripheral male Japanese macaques (Macaca fuscata) at Arashiyama West were studied during the 1994/1995 mating season. Although all peripheral males increased their proximity to the main troop, there was great variation in this behavior, from those who became virtually indistinguishable from the main troop males to those who moved in and out of the main troop in a clandestine fashion, to those who had only visual contact, from a distance of 25 m, with the main troop. The subsequent behavior of males displaying these patterns was compared, as was the behavior of peripheral to main troop males. Specifically, they were investigated for variation in three behavioral strategies which may function to increase access to mates: aggressive intimidation; affiliation with unrelated main troop females; and courtship display. The results show that peripheral males are a more diverse group than previously described, and that one group of peripheral males was able to successfully join the main troop, becoming indistinguishable from main troop males with regard to these behaviors which enhance access to mates.

Vigilance, predator detection and the presence of supernumerary males in vervet monkey troops

To determine whether adult males provide females with a service, in this case anti-predator behaviour, in return for inclusion in a group a free-ranging troop of vervet monkeys, Cercopithecus aethiops, was observed for 9 months. Males were more vigilant than females and the discrepancy increased as the breeding season approached. Higher levels of vigilance by males were associated with their spending a greater proportion of time either isolated or exposed at the tops of trees. Males, however, were not vigilant because they were exposed but moved into such positions in order to scan the surroundings. Despite this, males were no more efficient than females at detecting predators. The top-ranking male was the most vigilant and active against predators. It is argued from these and other data that male vigilance is directed at other males and is not a quid pro quo for inclusion. Although females benefit from this vigilance they are not capable of controlling male numbers, and troops, as a consequence, contain a number of supernumerary males who may well be the beneficiaries of the vigilance of others.

Life history variables of wild troops of formosan macaques (Macaca cyclopis) in Kenting, Taiwan

A study on population dynamics of wild Formosan macaques (Macaca cyclopis) in Kenting, southern Taiwan, was conducted from March 1985 to August 1990. At first, only one monkey troop was studied. It fissioned in 1987 and both of the daughter troops have been observed since then. Total number of animals increased from 10 to 29 over the years, but the sizes of individual troops have never been more than 20. Seasonality in breeding has been detected: copulations were concentrated in the period from November to January and 75% of all the 28 births occurred between April and June. Time of birth by parous females without offspring from the preceding year was earlier than that of lactating females. Young females gave their first births at 4 or 5 years of age. Total birth rate over the study period was 0.8 infant per female per year. Hunting was the main cause of death while natural mortality rate was low for the animals. There was only one adult male in each troop for most of the time. Troop males in the two daughter troops have been replaced two or three times in the three years by some solitary males that moved around in the area. The reproductive parameters of Formosan macaques in Kenting were found to be more similar to that of rhesus monkeys than to Japanese macaques. And a case of higher reproductive success in a high-ranking matriline was reported.

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.

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/km2 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.

Sexual strategies of female Japanese macaques (Macaca fuscata)

Evidence is reviewed that female Japanese macaques have multiple male mating partners when they are available and show a preference for mating with sexually unfamiliar males. Several lines of evidence suggest that this aspect of female sexual behavior results in the offspring of an individual female being sired by more than one male thereby maintaining the genetic diversity of the troop. Evidence is presented in this paper that a decrease in the number of adult troop males and a lack of extra-troop migrant males in the Arashiyama West troop of Japanese macaques following transplantation to a ranch in south Texas had consequences for the sexual behavior of the 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.

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.