Age-specific functions of stone handling, a solitary-object play behavior, in Japanese macaques (Macaca fuscata)

Stone handling (SH) in Japanese macaques, a form of solitary-object play, is newly acquired only by young individuals, and is the first example of a directly nonadaptive behavior that is maintained as a behavioral tradition within free-ranging provisioned social troops. We report here the first systematic investigation of this behavior in a stable captive social troop, the Takahama troop, which is housed in an outdoor enclosure of the Primate Research Institute (PRI), Kyoto University, Japan. This study was conducted to evaluate relevant competing hypotheses regarding the function of object play (e.g., misdirected foraging behavior and motor training) to explain the proximal causes and ultimate function(s) of SH. The "misdirected foraging behavior" hypothesis can be ruled out because of the lack of a clear temporal relationship between feeding and the occurrence of SH in any age class. Age-related differences in SH performance and behavioral patterns were observed, suggesting possible differences in the immediate cause and ultimate function between young and adults. Young individuals engaged in frequent bouts of short duration, involving locomotion and vigorous body actions throughout the day, which is typical for play by young in general. This pattern of behavior is consistent with the motor training hypothesis, which states that play occurs during the development of motor and perceptual skills and is thus potentially critical for neural and cognitive development. This practice is continued by those who acquire it at an early age, with adults engaging in significantly fewer but longer bouts that involve more stationary, complex manipulative patterns, almost exclusively in the late afternoon. We propose that for adults, at the proximate level SH is psychologically relaxing, but ultimately functions to maintain and regenerate neural pathways, and potentially helps to slow down the deterioration of cognitive function associated with advanced age in long-lived provisioned and captive macaques.     

Of stones and monkeys: testing ecological constraints on stone handling, a behavioral tradition in Japanese macaques

Japanese macaques are known to manipulate stones by displaying various seemingly functionless behavioral patterns, including carrying a stone, rubbing two stones together, or gathering several stones into a pile. This form of solitary object play called stone handling (SH) is a behavioral tradition in Japanese macaques, showing striking intertroop differences in frequency and form. Here, we evaluated two ecologically based hypotheses invoked to account for these differences. We hypothesized that the occurrence and form of SH would be affected by stone availability and the degree of terrestriality. We used standardized sampling methods to assess differences in SH and terrestriality among four captive and six free-ranging troops of Japanese macaques, and determine site-specific stone availability. Although we demonstrated that SH is almost exclusively a terrestrial activity, our comparative analyses showed that the number of stones readily available and the relative amount of time spent on the ground by the macaques were not associated with the intertroop differences in the occurrence of SH. Failure to accept the terrestriality and stone availability hypotheses suggests that the performance of SH and the motivation to engage in this activity are both more diverse and more complex than the direct links to time spent on the ground or the number of stones locally available. Other environmental influences and sociodemographic factors should be jointly considered to identify the sources of variation in SH, as a beginning to better understand the constraints on the appearance and subsequent diffusion of stone-use traditions in nonhuman primates. Am J Phys Anthropol, 2008. (c) 2007 Wiley-Liss, Inc.     

Environmental and Social Factors Associated with the Occurrence of Stone-Handling Behavior in a Captive Troop of <Emphasis Type="Italic">Macaca fuscata</Emphasis>

Stone-handling, a documented behavioral tradition in Japanese macaques (Macaca fuscata), occurs in both captive and provisioned free-ranging troops. We utilize data systematically collected as part of a broader investigation of stone-handling behavior in a captive troop to elucidate the environmental and social factors responsible for its occurrence. We analyzed contexts of stone-handling over 18 mo to determine under what conditions individuals most often perform it. There is clear seasonal variation in the occurrence of stone-handling. The lowest number and shortest duration of stone-handling bouts were in winter, gradually increasing to a peak in summer, and again decreasing toward autumn. Monkeys stone-handled more on clear sunny days than on cloudy or rainy ones. They displayed the behavior less under stressful conditions caused by human intervention or by aggression among troop members. Such stressful social conditions appeared to decrease individual motivation for stone-handling. In other words, individuals most frequently performed stone-handling under more relaxed environmental and social conditions. The findings are consistent with the hypothesis that stone-handling is a form of solitary object play behavior in macaques.     

Age-related differences in the performance, diffusion, and maintenance of stone handling, a behavioral tradition in Japanese macaques

Identifying the sources of behavioral diversity in non-human primates is vital to understanding the evolution of human behavior. Stone handling (SH, hereafter) is a form of object play consisting of the manipulation of stones by performing various behavioral patterns. This behavior is socially transmitted from generation to generation in Japanese macaques (Macaca fuscata), as a behavioral tradition. SH behavior in particular may reflect on the origin and evolution of stone-tool material culture. The objective of this study was to assess how group size, age structure, and age- and sex-related differences may account for the substantial intra- and inter-troop variations in SH reported in ten troops of Japanese macaques. Our results supported the hypothesis that patterns of variation in SH across troops reflected variability in group size and composition in age classes. We found that troop size was correlated with the proportion of troop members exhibiting SH simultaneously. The effect of troop size on the synchronized performance of SH may reveal the contagious nature of play. Our results suggest that the age structure of the group may affect the diffusion of SH. As predicted by the surplus energy hypothesis, a major functional hypothesis about play, intra-group variation in SH reflected more age- than sex-related differences. SH mainly occurred and was more frequent in younger than in older individuals, whereas no significant sex differences were found. SH episodes were shorter, more vigorous, and SH patterns were more diverse and less complex in immature than in mature individuals. The present findings reveal that age-related factors and group size may constrain the performance, diffusion, and maintenance of SH within a troop. Contrary to most other socially transmitted stone-tool using behaviors in non-human primates and early hominids, there is no optimal SH pattern. Provided some form of social learning, the non-adaptive nature of SH may allow particular SH pattern preferences to emerge at the group level. Our findings urge the use of an inter-populational comparative approach based on multivariate analyses when addressing the question of the evolution of behavioral traditions in primate and human populations.     

Food-snatching behavior of free-ranging Japanese macaques observed on Shodoshima Island: a preliminary report

We investigated the characteristics of a particular food-snatching behavior in which one individual forced another’s mouth open and grabbed the food, as performed by free-ranging Japanese macaques (Macaca fuscata) in Choshikei Monkey Park on Shodoshima Island, western Japan. We conducted a survey in late June 2012 and observed one of two monkey troops, comprising 214 monkeys. We recorded the age classes and sexes of the individuals who performed the snatching behavior and were snatched from, and examined the effects of provisioned food distribution and quantity on the frequency of snatching trials and success. During the survey, we recorded 747 snatching trials, of which 609 were successful, all of which were performed by seven individuals: one adult male and six adult females. The snatching behavior occurred only during provisioning. The target animals were primarily juveniles (650 trials, 578 successful), while cases in which food was snatched from adult females (93 trials, 30 successful) and subadult females (4 trials, 1 success) were less frequent. Among the juveniles, small juveniles had food snatched more frequently than large juveniles. The higher frequency of snatching trials against juveniles was likely due to their subordinate nature. Neither the distribution nor quantity of the provisioned foods had significant effects on the number of snatching trials and successes, while the time elapsed after provisioning had significant negative effects, attributed to a decrease in the number of wheat grains left within the mouth pouch of the potential target animals.     

History and Present Scope of Field Studies on <Emphasis Type="Italic">Macaca fuscata yakui</Emphasis> at Yakushima Island,Japan

Field studies on Japanese macaques on Yakushima Island started in the mid-1970s, >25 yr after the emergence of Japanese primatology, in response to criticism of methods using provisioning and the desire to find the socioecological factors influencing the social life of macaques in natural habitats. We habituated macaques without provisioning mainly in the coastal warm-temperate forest and found that they lived in small troops with a high socionomic sex ratio. Observations of several troop fissions and troop takeovers by nontroop males suggest that Yakushima macaques have a different social organization from that of Japanese macaques in other habitats. For example, youngest ascendancy as the dominance relationhip among sisters, which usually occurs in provisioned troops, was absent in Yakushima macaques. We compared their ecological and social features with those of Japanese macaques at Kinkazan (cool-temperate forests) and found that abundance of high-quality foods may cause stronger intra- and intertroop competition at Yakushima. Female Yakushima macaques may more positively solicit nontroop males to associate with them during the mating season. Such a tendency may promote frequent male movement between troops and frequent troop fissions. Though ecological factors form social features of Japanese macaques, some features such as male association and movements between troops are not accounted for via socioecology. Recent field studies have focused on macaques living at higher altitudes in Yakushima and on individual survival strategies by taking diverse viewpoints and using new technologies. DNA analysis of fecal samples shows low genetic diversity and suggests the macaques’ recent expansion from lowland to highland forests in Yakushima. The population censuses conducted annually indicate that the higher-altitude macaques have a larger home range but a similar group size versus their counterparts at low elevations. The unsolved issues in socioecology will pose a challenge to the younger generation of primatologists. Conservation of macaques and their habitat is one of our major activities at Yakushima. The level of protection has gradually increased in the National Park at Yakushima and, via our various conservation efforts, its most important area was designated a World Heritage Site by UNESCO in 1993. However, large-scale logging in the 1960s and 1970s caused the loss of macaque habitats and led to increased crop damage by them in the 1980s. We have proposed effective methods to protect cultivated fields from macaques as well as several plans for sustainable use of forests, such as ecotourism and a fieldwork course for university students. Local residents and researchers have created several nongovernment organizations (NGOs) to promote conservation and nature study at Yakushima. The role of local NGOs is particularly important to mitigate conflicts between people and wildlife. Though hundreds of macaques are still captured as pests annually in Yakushima, we continue the conservation measures and spread awareness of conservation in cooperation with the local NGOs.     

The Hand That Feeds the Monkey: Mutual Influence of Humans and Rhesus Macaques (<Emphasis Type="Italic">Macaca mulatta</Emphasis>) in the Context of Provisioning

Historically, humans and other primates (primates henceforth) have coexisted across cultures and contexts, and many primate populations use anthropogenic food sources as their main or supplementary food. While primates may actively forage for such food, they are also directly provisioned by humans in many regions. Ethnoprimatology views humans and primates as cohabitants of integrated socioecological spaces who mutually influence each other’s ecologies and social lives. We contextualized provisioning of primates by humans within an ethnoprimatological framework and examined if the availability of anthropogenic food affected primate diets or the amount of time primates spent in anthropogenic habitats and whether primates influenced the human act of provisioning. To this end, we used scan sampling on a group of rhesus macaques across a year, and conducted interviews with 86 people who paused at a nearby tea shop for refreshments. We found that the macaques’ consumption of natural resources and dietary diversity decreased, and they spent more time in human-modified habitats when provisioned food was available. We also found that particular behaviors of the provisioned macaques stimulated provisioning by humans. Our findings show that provisioning influences macaque feeding ecology and habitat use, and that the behavior of the macaques themselves drives people to provide them with food subsidies, illustrating a complex web of interactions between the sympatric species.     

Gut microbiota composition of Japanese macaques associates with extent of human encroachment

In recent decades, human–wildlife interaction and associated anthropogenic food provisioning has been increasing and becoming more severe due to fast population growth and urban development. Noting the role of the gut microbiome in host physiology like nutrition and health, it is thus essential to understand how human–wildlife interactions and availability of anthropogenic food in habitats can affect an animal's gut microbiome. This study, therefore, set out to examine the gut microbiota of Japanese macaques (Macaca fuscata) with varying accessibility to anthropogenic food and the possibility of using gut microbiota as indicator for macaques’ reliance on anthropogenic food. Using 16S ribosomal RNA gene sequencing, we described the microbial composition of Japanese macaques experiencing different types of human disturbance and anthropogenic food availability—captive, provisioned, crop‐raiding, and wild. In terms of alpha diversity, our results showed that observed richness of gut microbiota did not differ significantly between disturbance types but among collection sites, whereas Shannon diversity index differed by both disturbance types and sites. In terms of beta diversity, captive populations harbored the most distinctive gut microbial composition, and had the greatest difference compared with wild populations. Whereas for provisioned and crop‐raiding groups, the macaques exhibited intermediate microbiota between wild and captive. We identified several potential bacterial taxa at different taxonomic ranks whose abundance potentially could help in assessing macaques’ accessibility to anthropogenic food. This study revealed the flexibility of the gut microbiome of Japanese macaques and provided possible indices based on the gut microbiome profile in assessing macaques’ accessibility to/reliance on anthropogenic foods.     

Comparison of stone handling behavior in two macaque species: implications for the role of phylogeny and environment in primate cultural variation

This study describes the stone handling (SH) behavior of captive rhesus macaques and compares it with that of a captive troop of Japanese macaques with reference to the relative contributions of phylogeny-driven behavioral propensities, environmental differences and socially facilitated learning to the formation of culture. These systematically collected data demonstrate for the first time that two closely related macaque species might share a common cultural behavior, SH. The rhesus troop displayed SH behavioral patterns that was already described in Japanese macaque troops. The one exception was a new pattern not yet seen in any Japanese macaque troop. Differences in the physical environment of the two study enclosures may be responsible for some of the variation in observed SH behavioral patterns in these two troops. These data support the idea that environmental factors can be important for the formation of cultural variation, when the key materials needed to perform the behavior are present in both habitats (stones). Our results are consistent with the prediction made by Huffman and Hirata [The biology of tradition: Models and evidence. Cambridge: Cambridge University Press. p 267-296] that an interactive triad of phylogenetic, environmental and social factors can be responsible for the formation of cultural variation in primates.     

Congenital malformations in Japanese macaques (Macaca fuscata) at Takasakiyama

From the late 1960s to the early 1970s, many congenitally malformed infants were born into provisioned Japanese macaque troops. Although the exact cause of this problem was not determined, the occurrence of malformations decreased thereafter. We examined possible factors such as total population size, number of adult females, birth rate, and volume of provisioned food. Agrichemicals attached to provisioned food are suspected as the main cause, as other factors were found to have no influence. Many more malformations were seen in males compared with females, in feet compared with hands, and in the fourth compared with other digits. We confirmed that the frequency of congenital malformation was high during the 1960s through to the mid-1970s when increased levels of provisioned food were given and that the incidence of congenital malformations was also elevated among wild macaques during this time.     

Primates,Provisioning and Plants: Impacts of Human Cultural Behaviours on Primate Ecological Functions

Human provisioning of wildlife with food is a widespread global practice that occurs in multiple socio-cultural circumstances. Provisioning may indirectly alter ecosystem functioning through changes in the eco-ethology of animals, but few studies have quantified this aspect. Provisioning of primates by humans is known to impact their activity budgets, diets and ranging patterns. Primates are also keystone species in tropical forests through their role as seed dispersers; yet there is no information on how provisioning might affect primate ecological functions. The rhesus macaque is a major human-commensal species but is also an important seed disperser in the wild. In this study, we investigated the potential impacts of provisioning on the role of rhesus macaques as seed dispersers in the Buxa Tiger Reserve, India. We studied a troop of macaques which were provisioned for a part of the year and were dependent on natural resources for the rest. We observed feeding behaviour, seed handling techniques and ranging patterns of the macaques and monitored availability of wild fruits. Irrespective of fruit availability, frugivory and seed dispersal activities decreased when the macaques were provisioned. Provisioned macaques also had shortened daily ranges implying shorter dispersal distances. Finally, during provisioning periods, seeds were deposited on tarmac roads that were unconducive for germination. Provisioning promotes human-primate conflict, as commensal primates are often involved in aggressive encounters with humans over resources, leading to negative consequences for both parties involved. Preventing or curbing provisioning is not an easy task as feeding wild animals is a socio-cultural tradition across much of South and South-East Asia, including India. We recommend the initiation of literacy programmes that educate lay citizens about the ill-effects of provisioning and strongly caution them against the practice.     

Non‐territorial Macaques Can Range Like Territorial Gibbons When Partially Provisioned With Food

Human food supplementation can affect components of animal socioecology by altering the abundance and distribution of available food. We studied the effect of food supplementation by comparing the ranging patterns and intergroup interactions of two groups of northern pigtailed macaques (Macaca leonina), a non‐territorial primate species. One group was partially reliant on food provisioning, whereas the other group foraged wild food. We also compared the macaques’ movement with that of a group of white‐handed gibbons (Hylobates lar), a territorial species inhabiting the same site. Home range, core area, and daily path lengths were significantly smaller for the semi‐provisioned group than for the wild‐feeding group. In contrast to wild‐feeding macaques, supplemented macaques showed higher fidelity to home range, core area, and particularly to the region where human food was most accessible and abundant. The relationship of daily path length and home range indicated a low defendability index for wild‐feeding macaques; the higher index for the semi‐provisioned group was consistent with the territorial pattern found in gibbons. Semi‐provisioned macaques showed further traits of territoriality with aggression during intergroup encounters. These findings indicate that human modification of food availability can significantly affect movement patterns and intergroup competition in macaques. The observed ranging dynamics related to food provisioning may decrease the efficiency of macaques as seed dispersers and increase predation on their home range, and thus have important consequences for plant regeneration and animal diversity.     

Changes in Demographic Parameters of <Emphasis Type="Italic">Macaca fuscata</Emphasis> at Takasakiyama in Relation to Decrease of Provisioned Foods

Though researchers have studied population dynamics extensively in many free-ranging primate populations, information on the relationship between food abundance and demographic parameters based on long-term data is sparse. We examined changes in demographic parameters in relation to decreased provisioning of foods based on data gathered for >50 yr in a provisioned, free-ranging Japanese macaque (Macaca fuscata) population at Takasakiyama, Oita Prefecture, in the southern region of Japan. At Takasakiyama, population size increased linearly because of heavy provisioning in the 1950s and 1960s. The provisioning of food to macaques decreased from 1965. We examined changes in the physique index of full adult females, primiparous age, birth percentage, infant mortality rate, population density, and annual population growth rate relative to the decrease of provisioned foods between the 1970s and 1990s. As a result of the drastic decrease in the amount of foods provisioned, the physique index of full adult females decreased, primiparous age increased, birth percentage decreased, and infant mortality rate increased, such that annual population growth rate diminished substantially. Ours is the first study to demonstrate quantitatively the relationship between provisioned food abundance and demographic parameters based on long-term data.     

Symmetrical patterns in non-agonistic social interactions found in unprovisioned Japanese macaques

Non-agonistic social interactions in an unprovisioned troop of Japanese macaques (Macaca fuscata yakui) were analyzed with the spacing between individuals, leading-following interactions, and exchange of social grooming. The most frequent interactions were found between kin-related females. Unrelated females stayed with one another rather frequently, but rarely exchanged social behaviors. Interactions between males and females were infrequent though they were occasionaly observed between high-ranking males and high-ranking females. Very frequent exchange of grooming was observed between males, and even high-ranking males exchanged grooming more frequently with males than with females. Most non-agonistic social interactions in the study troop were based on bidirectional exchange of social behaviors, in which no clear tendency relevant to dominance or sex was found; while in provisioned Japanese macaque troops, associations between males and females, between unrelated females, and between males were formed mainly be subordinates' active roles in associative behaviors. This seems relevant to the idea that dominance grealty influence social life in provisioned troops. The present study provides guidelines for interspecific comparison of social interaction patterns of macaque species.     

Why dominants do not consistently attain high mating and reproductive success: A review of longitudinal Japanese macaque studies

There is wide interest in the effects of reproductive biology, mating partner preference, and rank on mating success (MS) and reproductive success (RS) in primates. In particular, theory stresses importance on the mechanisms for attaining RS. Most theory hedges on competitive ability and priority of access to resources, whether they be food or estrous females. However, the majority of data used in favor of such hypotheses come from relatively short-term studies. We review these hypotheses based on long-term data from provisioned and unprovisioned populations of Japanese macaques. Neither MS nor RS were consistently attained by high-ranking males and females. For males, female choice and mating partner preference is seen to over-ride most male-male competitive behaviors likely to affect MS and RS through priority of access to estrous females. Long-term mating patterns driven largely by female partner preferences, results in decreasing MS and RS for older higher-ranking males. The long-term trend for females to prefer less familiar or novel partners results in higher MS and RS for younger, middle-ranking males. The effects of this vary according to troop size and the duration of male tenure. For females, no consistent trend was recognized for rank related RS in either provisioned or unprovisioned troops. Non-reproductive mating may provide differential benefit to high-ranking females for access to limited food resources in some habitats but overall the relationship was inconclusive. Distribution and defendability of food resource, rather than provisioning per se may be more important.     

Stone handling behavior in rhesus macaques (Macaca mulatta), a behavioral propensity for solitary object play shared with Japanese macaques

Stone handling (SH) behavior was systematically studied in a captive troop of rhesus macaques housed at the Primate Research Institute of Kyoto University, and compared with the results of long-term studies of this behavior in Japanese macaques, to evaluate the similarities of SH behavior in these two closely related species. Similar to Japanese macaques, rhesus macaques showed age-related differences in SH. Young animals were more active and displayed more SH patterns and bouts than did adults. Furthermore, the young displayed SH at a higher frequency and their bouts were of a shorter duration, compared to adults. Young adults were more active and displayed more patterns than did older adults. On the other hand, older adults were more conservative and displayed fewer patterns, and engaged in them for longer durations. All individuals displayed SH more frequently in relaxed environmental and social conditions. While lacking an apparent immediate adaptive value, practice of the behavior has been proposed to have long-term functional value for neural and cognitive development in the young and for the maintenance or repair of neuro-pathways in aging macaques that habitually perform the behavior. The results presented here are consistent with what we know about Japanese macaque SH. Given the uniformity of SH behavioral parameters and these two macaque species’ close phylogenetic relatedness, we propose that a similar functional and adaptive value for SH can be inferred for rhesus macaques.     

Status of urban populations of the long-tailed macaque (<Emphasis Type="Italic">Macaca fascicularis</Emphasis>) in West Sumatra,Indonesia

We studied long-tailed macaque (Macaca fascicularis) populations in Padang, West Sumatra, Indonesia, focusing on the effect of human provisioning on their demography and dietary composition. We conducted a field survey at three sites in the city: Gunung Meru, Gunung Padang, and Gunung Panggilun. Mean troop size (range 28–68) and infant ratio (range 0.38–1.00) were greater in Gunung Meru, where the macaques have been highly provisioned, than at the other two study sites (troop size 10–15; infant ratio 0.00–0.33). The macaques at all sites consumed both natural and human foods, but dependence on the latter differed among sites: three-quarters of the diet of macaques in Gunung Meru consisted of human foods, while human foods comprised less than 5% of the macaque diet at the other sites. The ability of macaques to modify the proportion of human food is a behavioral flexibility that facilitates the survival of the long-tailed macaque in urban habitats. Without restrictions on provisioning, the degree of dependence of macaques on human foods and population size could increase, especially in Gunung Meru, and human–macaque conflict could escalate. In order to create an effective management policy for urbanized monkeys, long-term quantitative data on macaque behavior and monitoring of population parameters are required.     

人工投食行为对非人灵长类社会生态的影响

近年来,国内开展了对几种非人灵长类投食群的研究。借鉴日本猴(Macaca fuscata)、恒河猴(M.mulatta)和黑猩猩(Pan troglodytes)长期积累的研究经验,有利于我们减少人工投食对非人灵长类带来的不利影响,也有利于推动国内非人灵长类社会生态学的深入研究。本文通过比较这三种灵长类野生群和投食群之间的差异,量化地分析了人工投食对灵长类动物社会生态学方面的影响。结果表明在社会行为方面,人工投食会导致灵长类动物的聚集、增加争斗频率、改变个体间等级关系、延缓分群现象、影响活动域和增加新兴行为。在种群数量方面,人工投食会缩短雌性性成熟时间、增加出生率和婴猴存活率、减少成年猴死亡率。而人工投食对雄性性成熟、出生间隔等方面的影响仍需要进一步考证。最后我们讨论了人工投食行为的利弊,并建议应谨慎和妥善地实施人工投食,和对非人灵长类投食群进行合理的监管。     

Probability of paternity exclusion and the number of loci needed to determine the fathers in a troop of macaques

We calculated the probability of paternity exclusion (P) in 6 troops of rhesus and Japanese macaques housed in open enclosures and 68 wild troops of Japanese, crab-eating, and toque macaques using 33 genetic loci which encoded the blood protein variations detected by electrophoretic techniques. In the open enclosures, especially of rhesus troops, we obtained a fairly high probability of paternity exclusion and succeeded in determining the fathers of offspring. However, we found significant differences between the observed and calculated probabilities in most of the troops. These differences were ascribed to a situation whereby the Hardy-Weinberg equilibrium had not been attained in the troops and/or the numbers of variable loci were too small. In the wild troops of Japanese, crab-eating, and toque macaques, the means ofP were 0.2274 (0.0192–0.5017), 0.4635 (0.1676–0.7151), and 0.7382 (0.6266–0.7954), respectively. We also estimated the number of loci needed to determine the fathers of offspring with a probability of 0.8 assuming that ten males were present in the troop. The estimated number was about 13.5 times, 5 times and 1.8 times the number of loci examined on average in the troops of Japanese, crab-eating and toque macaques, respectively. This means that determination of most of the fathers of offspring in wild troops of these macaques, even of toque macaques which have a rather high probability of paternity exclusion, is difficult so long as we employ only electrophoretic techniques.