Intra-troop spacing mechanism of the wild Japanese monkeys of the Koshima troop

The average frequencies of communicative behavior, social behavior, and social encounters (inter-individual proximity within three meters) per hour for a monkey were obtained in their natural habitat by tracing several adult males and females of a Japanese monkey troop living in the Koshima islet. The spatial distribution patterns and the density of troop members within the expanse of the troop at any moment were investigated by tracing several adult femals. Frequency distributions of the monkeys found within five and 10 meters were compared with a Poisson distribution. The frequencies of social encounters and of social interactions of Japanese monkeys were distinctly low, except between mothers and their offspring. The density of monkeys within the expanse of the troop at any moment was very low. Both aggressive behavior and inter-individual proximity (within three meters) were distinctly low when monkeys were foraging for natural food. An avoiding mechanism among troop members plays an important role in maintaining the social structure of these Japanese monkeys. This mechanism works in two ways: each individual does not approach others too closely; the density of monkeys within the expanse of the troop is low at all times.     

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.     

Parasite changes and their influence on the body weight of Japanese monkeys (Macaca fuscata fuscata) of the Koshima troop

The incidence of eggs in individual feces (percentage of infected animals) and the EPG (eggs per gram of feces) of gastrointestinal nematodes for individually discriminated Japanese monkeys of the Koshima troop were investigated monthly from October 1974 to June 1979. Eggs of four nematode species,Oesophagostomum aculeatum, Trichuris trichiura, Streptopharagus pigmentatus andStrongyloides fülleborni, were frequently found; a few cestode segments ofBertiella spp. were occasionally found.S. fülleborni had a high incidence (100%) in young monkeys, suggesting that its incidence might vary with the monkeys' age. Seasonal changes in the incidence and EPG of each parasite were observed. The monkeys were treated with the anthelmintic thiabendazole once a month from May to July, 1975. Thiabendazole exerted an excellent effect onS. pigmentatus andS. fülleborni, and a lesser effect onO. aculeatum. Decreases in parasites caused by either seasonal changes and/or the anthelmintic treatment did not produce an increase in the monkeys' body weight, indicating that body weight is more affected by food intake or some factors other than parasitic infections. It is suggested that field anthelmintic treatment on wild monkeys should be conducted with the greatest care, since reinfection can occur immediately after such treatment.     

Retardation of bone development in the Koshima troop of Japanese macaques

Retardation of bone development was observed in the Koshima troop of free ranging Japanese macaques. In the control group, epiphyseal unions of appendicular long bones generally started to close at about 4 yrs of age and were completed at about 8 or 9 yrs of age. Limb bone unions of the Koshima troop, however, started to close at about 9 yrs of age and completely closed at about 15 yrs of age. In the epiphyseal unions of trunk and girdle bones, the Koshima troop again showed a retardation of closure compared with the control group. Until long bones reached their full length, that is, until about 15 yrs of age, their size was small in the Koshima troop compared with the control group, though the sample size of the Koshima troop was small. After 15 yrs of age, however, many osteometrical measurements of the Koshima troop were nearly the same as controls. A prolonged growing duration compensated for the slow growth and allowed them to become as large as controls. This prolongation may be an adaptation in response to small size during the developmental period. In some parts of the body, however, Koshima macaques failed to reach the adult size of controls. Males were less likely than females to reach full size. Causes of the retardation and small size in the Koshima troop are discussed, but they remain open to further studies.     

The social interactions of juvenile Japanese monkeys on Koshima Islet

This study reports on the social interactions of juvenile Japanese monkeys (1–5 years old) living in their natural habitat. Juvenile males preferred to associate and play with males of the same age. Male-male interactions initiated by a subordinate of the same age or by an older frequently involved play. Juvenile females tended to interact with younger individuals, and to avoid older males. Older females (4–5 years old) tended to avoid one another. Sibling pairs had high frequencies of interaction, although the occurrences of particular types of interactions varied from pair to pair. Patterns of social relations among adults are discussed in relation to juvenile behavior.     

Food and nutritional condition of free ranging Japanese monkeys on Koshima Islet during winter

A female with infant was chosen as material for study in an attempt to assess the nutritional condition of free ranging Japanese monkeys during winter. Her daily food composition, dry weight intake and nutritional (protein, lipid, carbohydrate, ash and calorie) intake were measured monthly (October to March). About 90% and 8% of the autumn diet consisted of fruits and invertebrate animals, respectively, while 70% of the winter diet (February) consisted of leaves of evergreen trees. Comparing the daily protein intake of this focal female with the requirement level estimated from references, only October and November represented months fulfilling this level. Also, there was a remarkable decrease in lipid intake towards winter. These results coincided well with the observed body weight loss in the female and the increasing feeding activity of her baby towards winter. It is suggested that such seasonal malnutrition of the mother might affect population parameters such as the infant mortality.     

Activity patterns of a troop of Japanese monkeys (Macaca fuscata yakui) on Yakushima Island,Japan

Activity patterns of Japanese monkeys (Macaca fuscata) were observed for 240 hr from August to December 1976 on the western slope of Mt. Kuniwari, Yakushima Island, Japan. Activity patterns and the time budget of a habituated wild troop which consisted of 47 animals in August 1976, were studied quantitatively by using the scan-sampling method at 15-min intervals. Six thousand seven hundred and six animals were recorded in 959 scans during the study period and the mean number of animals seen per scan was 7.0. The time budget established for different categories of activity was as follows: inactive—20.9%; moving—22.8%; feeding—23.5%; social grooming—27.9%; self-grooming—1.2%; and other activities—3.7%. Adult males spent less time in feeding and more time in resting or being inactive than females or juveniles. The daily activity patterns were highly variable with respect to time. Intraspecific variations were examined between troops in several regions of Japan and it was noted that the percentages of time devoted to feeding were similar in all areas. Inter-species variations in the activity budgets of several species of primates were also examined. The percentage of time spent in social grooming by Japanese monkeys is exceptionally high compared to that recorded in other species.     

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

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

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.     

Fish: a new addition to the diet of Japanese macaques on Koshima Island

Recently, Japanese macaques on Koshima Island have begun to eat raw fish. This new habit was first observed in 1979, in adult males that live on the periphery of the group of macaques. The habit was first transmitted to the old dominant females in the group and then spread gradually to most other members of the group over the next 6 years. However, Koshima macaques do not appear to have a particular liking for fish. The eating of fish is discussed in relation to recent nutritional conditions in the macaques' habitat.     

The reproductive biology of a free-ranging troop of Japanese monkeys

Quantitative data on the reproductive biology of Japanese monkeys (Macaca fuscata), especially of females, are reported here. Arashiyama B troop shows stable annual breeding periodicity, and has mating seasons between October and March. In mating seasons, females show estrus with rather consistent cyclicity, and are in estrus for a total of 35 days on an average. The mean length of estrous cycle is 34 days. Although most females show estrus inter-menstrually, some show peri-menstrually; i.e., estrus does not always synchronize strictly with ovulation. Pubescent females between 3.5 and 6.5 years show estrus, but with low sexual activity. Middle-aged females between 7.5 and 16.5 years show high sexual activity with high conception rate. Old-aged females over 17.5 years show low sexual activity. Females with surviving infants have lower sexual activity than those without infants. There is found no significant correlation between female's ranking and sexual activity. This study was financed partly by the Cooperative Research Fund of the Primate Research Institute, Kyoto University.     

Longitudinal changes of dominance rank among the females of the Koshima group of Japanese monkeys

The changes of dominance rank among female Japanese monkeys of the Koshima group over a period of 29 years from 1957 were studied. The dominance rank order was relatively stable in the early population growing phase, while large scale-changes of dominance rank order occurred successively in the phase of population decrease brought about by the severe control of artificial feeding after 1972. Nevertheless, the rank order of several females of the highest status was stable. Furthermore, the reproductive success of these highest status females was high (Mori, 1979a;Watanabe et al., in prep.). Divergence of the dominance rank order fromKawamura's rules (Kawamura, 1958) was observed in the following respects: (1) Some females significantly elevated their rank depending on the leader males. (2) If mothers died when their daughters were still juveniles or nulliparous, the dominance rank of some of these offspring females was significantly lower than the mother's one. However 55% of daughters which lost their mothers at a young age inherited the mother's rank. (3) Dominance among sisters whose mother had died when at least one of the daughters was under 6 years old followed the rule of youngest ascendancy in 60% (Kawamura, 1958), and in 80% when both of the daughters were nulliparous at the mother's death. The mean rate of aggressive interactions for each female with subordinates to her was calculated by dividing the total aggressive interactions between the female in question and her subordinates by the number of subordinate females to the female in question. A female which showed a high rate of aggressive interactions with her subordinates was categorized as an “Attacker”, and a female showing a lower rate was categorized as a “Non-attacker”. Similarly, categories of “Attacked”, and “Non-attacked” were distinguished by using the rate of aggressive interactions with dominant females. Several females which were once categorized in one category in a year were repeatedly categorized in the same category over different years. The “Attacked” tended to be females of higher rank, and “Non-attackers” tended to be females of lower rank. “The second-higher-status females”, were “Attacked”, and their rank was unstable. In particular, females of lower rank within the lineage of the highest rank suffered this kind of severe status. Most of the daughters of these females showed a sharp drop of rank, and died when they were still at a young age, i.e. “the second-higher-status females” displayed low fitness. “Non-attackers” were significantly “Non-attacked”; i.e. they were females which showed a non-social attitude. Females which underwent a drop of rank tended to be “Non-attackers”. The most important factor which determined the females' rank was the memory of their dominance relations under the influence of their mother [dependent rank (Kawai, 1958)] in their early life during development. This finding corresponds well with the results in baboons obtained byWalter (1980); the target females of aggressive interactions by adolescent females were determined by the rank of the mothers when these adolescent females were born.     

Maternal and alloparental care in a troop of free-ranging Japanese monkeys

In four years of observation of a free-ranging Japanese monkeys, reproductive rate, physical development of infants, maternal care and alloparental care were studied. Primiparous mothers, orphanned mothers and old mothers showed different types of maternal care from typical multiparous mothers, and the reproductive success of each of these categories of mother also differed. Among nulliparae, 2-year-old females showed the most interest in infants. Adult females were generally indifferent to other females' infants and came to act aggressively toward them with increasing age and maternal experience.