In: The Monkeys of Arashiyama Thitty-five years of research inJapan and the West. (eds.) Linda M. Fedigan & Pamela J. Asqwith. (1991) State University of New York Press. 1 History of the Arashiyama Japanese Macaques in Kyoto, Japan Michael A. Huffman INTRODUCTION A history of research at Arashiyama is incomplete without a brief outline of the beginnings of primatology in Japan (also see Itani 1983, 1985a). In 1948, at Toimisaki, Miyazaki Prefecture, Kyushu (fig. 1), Kinji Imanishi and two of his students, Shunzo Kawamura and Junichiro Itani, started a sociological study of semiwild horses. Influenced by Darling’s work on red deer (Darling 1937), Imanishi had planned to observe mammals by the use of individual identification (Imanishi 1955). At the beginning of December, while at Toimisaki, they decided to travel 12 kilometers up the coast to the small island Koshima where they had heard from local people that there were wild monkeys. The first visit on December 3rd was unsuccessful be- cause they could not find the monkeys; however, after returning to Toimis- aki they occasionally saw monkeys there (Itani 1985a). The Primates Research Group was, in practice, operating from 1948 with the advent of Imanishi, Kawamura, and Itani’s studies. However, ac- cording to Denzaburo Miyadi (1966), the group was officially formed in November of 1951 at Kyoto University’s Laboratory of Ecology by Imanishi and Miyadi. Included in the group were Itani, Kawamura, Naonosuke Hazama, Masao Kawai, Tadao Umesao, and Kisaburo Tokuda. The group conducted a series of preliminary surveys of the Japanese macaque habitat throughout the country, from Yakushima at the southern- most boundary of the species distribution, to Shimokita, the northernmost boundary, including such places as Takasakiyama, Minoo, Arashiyama, Hieizan, Shodoshima, Boso, Tsugaru, and Towada (fig. 1). Toimisaki, Ko- shima, and Takasakiyama were chosen as the first three study sites. Takasakiyama was first visited in April of 1950 by Kawamura and Itani (Itani 1961, 1985a). 21‘WLIIOWIHS "TI fvonunsl “11 ‘vaVMOL “OI {(VWVAODVAV) OSOg °6 ‘NVZISI °g ‘VAVAIHSVay “L ‘OONIW “9 SVWIHSOGOHS °S SVAVAINVSVVL “VY SVWIHSOY “€ SIIVSINIO] *7 SVWIHSAAY JX “I SALISHAAIN(| OLOAM x “NVZISIH GNV VWVAIHSVEy ‘SALIS VIUV OLOAY OML FHL JO dVW GATIVLGG HLIM NVdVf NI SALIS AGNIS ATANOW JSaNVdVE “TL AINOLYHistory of the Arashiyama Japanese Macaques 23 For the first few years, it was quite difficult to observe the monkeys at these sites. The best the researchers could do was to catch glimpses of them as they traveled along exposed parts of the mountainside or while up in the trees feeding (Itani 1954). Because Japanese monkeys had been a hunted species for many years, they were afraid of humans. In order to observe the monkeys for longer periods of time, attempts were made at Koshima to provision them along their travel routes in the forest, and these efforts proved successful. Kawamura and Itani originally developed the idea of provisioning the monkeys from a local farmer, Toui- chi Kanchi, who told them that he had fed the monkeys sweet potatoes some years earlier (Itani and Tokuda 1958; Itani 1985b). In 1951 and 1952, Imanishi, Kawamura, Itani, and Tokuda carried out a series of investigations on Koshima (Itani 1985a). By August of 1952, Itani and Tokuda were able to observe the group out in the open as they came to the beach to feed on provisioned sweet potatoes (Itani and Tokuda 1958). During this same period the monkeys at Takasakiyama were also being provisioned by Shino Onishi, a priest from the temple at the foot of the mountain for the purpose of preventing damage to local crops by the monkeys and for tourism (Itani 1985a). The Primates Research Group focused its attention mainly on the Ko- shima and Takasakiyama groups. From this point, long-term studies based on individual identification were conducted, resulting in pioneering work on dominance rank order (Kawai 1958a, 1958b; Kawamura 1958), parental care (Itani 1959), cultural behavior (Kawamura 1958; Kawai 1965), vocal communication (Itani 1963), and sexual behavior (Tokuda 1961-62). THE EARLY YEARS AT ARASHIYAMA The earliest account of any observations of the Japanese macaques at Arashiyama is given by Eiji Ohta, a naturalist and now retired primary School principle in Kyoto. Ohta (1975) says that he first encountered the troop in 1948 by accident while looking for tree frogs. Thereafter, he occa- sionally attempted to follow and observe the group. In 1951, Itani and Kawamura made a brief survey of the Arashiyama area but were unable to directly observe the monkeys. From indirect evidence obtained during the survey, they estimated that a group of around 50 individuals inhabited the area (Hazama 1972; Asaba 1984). The first long-term study of the group was conducted by Hazama, then affiliated with Kyoto University, and it is to him and his early coworkers that we owe much for the documentation of the Arashiyama troop’s early history. According to Hazama (1972), the first plans to study and provision the Arashiyama group were made on July 14, 1954, by two entrepreneurs of the Arashiyama area, Sonosuke Iwata and Eiji Furukawa. A meeting was24 Huffman held at Furukawa’s Inn (Togetsutei) close to the present entrance of the park. Ten people were present including Furukawa, Iwata, Hazama, Miy- adi, a representative of the Kyoto Zoo, the head of the Department of Ag- riculture and Forestry of Kyoto city, and a representative from the Kyoto Tourist Commission. It was stipulated that the use of the national forest as the final provi- sioning site and park grounds was to be avoided. Since financial support from local government agencies was prohibited, it was agreed that Iwata would privately manage the park, providing .15 square kilometers of his own land. Hazama was put in charge of the preliminary studies and of lur- ing the monkey group to the proposed site at Iwatayama. He started his investigation on October Sth of that year (Hazama 1972). In the beginning, Hazama (1972) found that locating the monkeys was very difficult. At the time, the group’s range was approximately eight square kilometers; extending from Arashiyama to the Kamome valley (fig. 2). Apparently, the group stayed in the national forest of Arashiyama only during late autumn and early winter to feed on the fruits of Aphanathe aspera and Celtis sinensis. During the rest of the year they stayed in Ka- mome valley (Hazama 1972). Occasionally accompanied by Furukawa, Hazama spent days walking through the forest getting acquainted with the animal trails that criss- crossed the rugged terrain. With little success in locating the group, Hazama started asking people in the area about where and when monkeys had been seen. Although Hazama found the local hunters to be a good source of information they were also a great hindrance. Hazama writes: I was beginning to lose hope, wondering if the monkeys had been chased out of the area. . . . Upon second thought, however, I reconsidered. De- spite the fact that Kyoto is the center of Japanese primatology, this area was relatively untouched due to the difficulty in locating the monkeys. I was not going to give up so easily. (1972) In 1955, the second year into Hazama’s study, conditions became fa- vorable for provisioning, as the group’s habitat in Kamome valley (fig. 2) was drastically affected by the cutting down of the forest. The area from Karasugatake to Hatogasuyama was timber-mined. Research progress was still slow, but the men were able to catch glimpses of a few monkeys. Around January of 1955 Hazama and his assistants experimentally began to set out sweet potatoes in the national forest (Hazama 1972). At first, the food went untouched, but in February and March, one of the leanest periods of the year for the monkeys, potatoes and other seasonal fruits scattered about the area would disappear within a few days. Hazama and Furukawa, with the help of Shizuma Hirose, Kinya Nakajima, and others,“I ‘Ola ‘9961 ‘bz “Dld ‘7961 VNVZVH WoOud GaLldvay “THAR VES AAOMV SAMLAW NI SHGNLILIV GALVNOISA “SNIWHS INVOAAVY “C ‘IONVALNI WVg °O ‘NOLLVLS HOUVaS -FY AYOLSI TWANLVN VWVAIHSVYY GNV WNASNY “G !NOILVLS LNIWFOVNVW-YOLISIA GNV ALIS ONINOISIAON “WV Kem ite et pane Wyed3o0} POUBPIM “PLOY mm 210}0q door} eweAwyseay yo e8uel ewOH >) “AUVd ADANOW VAVAVLVM] HL NI aLIS ONIGHa LNAYAND FHL ANY ‘ONINOISIAONd AOI dNOUD VNVAIHSVAY AHL JO ALNOU OIGVWON ONY ZONVY JWOH ‘t BUNS,26 Huffman took advantage of this and established their first temporary feeding site (Hazama 1962; Hirose 1984). They all wore white chest aprons with a large red ball painted in the middle, symbolic of the Japanese flag. This was done in an attempt to make the monkeys familiar with their presence, in the hope that by associating this symbol with provisioned food, the monkeys would differentiate these men from the hunters (Hirose 1984). The feeding site was moved about 100 meters at a time, moving along the route that the monkeys naturally followed from Kamome valley to the national forest (fig. 2; Hazama 1972). At the second feeding site, a male (later named Shan) came up to Hazama for the first time and picked up an apple that Hazama had gently rolled onto the ground. The group grew more used to people and at the third feeding site, Hazama counted 47 individuals. However, calculating back from Hazama’s records after provisioning, it is estimated that in the fall of 1954, the troop consisted of 34 individuals; 20 females and 14 males, plus an additional seven identified solitary males (Hazama 1962; Koyama 1984). On March 26, the seventh site was placed on their return route to Kamome valley. The group returned and stayed in Kamome valley for the next six months, a nomadic pattern they continued until the third year of provisioning (Hazama 1962). On December 24, 1955, the last site was es- tablished on Iwatayama, the present location of the park’s provisioning sta- tion (Hazama 1972). Now that the entire group could be observed out in the open, Hazama, Nakajima, and Hirose began to identify its members. Interestingly, they as- signed names according to recognized personality traits or physical appear- ance (Nakajima 1964, 1984), rather than just a number or randomly selected name of an animal or plant, as was done at Koshima and at other sites in Japan and abroad. For instance, Nakajima named one female Coo- per, as she reminded him of the actor Gary Cooper. The names of the adult females were retained and later used as lineage names in the system begun by Nakajima, in which the year of birth is placed after the individual’s mother’s name (Koyama 1967). For example, Mino-6369? was born in 1969 to Mino-63@ and Mino-632 to Mino in 1963, making Mino the grandmother to Mino-6369¢. With this system the genealogical relationship and degree of consanguinity between two related individuals can be determined at an instant. SOCIAL CHANGE Since observations were begun by Hazama in 1955, the group has under- gone many changes. Two important elements of this social process have been group fission, and changes in alpha male and in alpha female status. To the present (1989), If males and nine females have been recognized to attain the status of alpha male and female, and four fissions have occurred.History of the Arashiyama Japanese Macaques 27 While changes in male and female alpha status were not necessarily depen- dent upon group fission, fission always coincided with changes in male and female status. This series of events is illustrated in figure 3. Male Emigration, Immigration, and the Death of a Matriarch Including Ohta’s independently conducted observations from 1948, this his- tory begins with a male named Mars, whom Ohta (1975) identified as the alpha male. He named the beta male Minerva and later stated that this was the same individual Hazama renamed Zao. By 1955, Zao had become alpha male, but disappeared in June 1959 (Hazama 1972; Ohta 1975). Between 1959 and 1964, Lincoln, Shan, and Y consecutively became alpha males of the troop. After each male disappeared he was replaced by the next (Hazama 1972; fig. 3). During this early period of primatology, it was thought that the Japanese macaque troop was a closed society (Itani 1954; Mizuhara 1957) and thus Hazama (1972) originally suspected that all these males had been killed by hunters. Hazama (1972) noted that during the period shortly after Zao’s disap- pearance, the alpha female, Tokiwa, became the highest-ranking individual in the troop. However over time, the subleader male Lincoln moved into the center of the group and became the next alpha male. After only 15 months as alpha male, he disappeared. Shan, described by Hazama (1972) as not being very aggressive, became the next alpha male. Norikoshi and Koyama (1975) later recognized that age and the length of stay in the group were correlated with dominance rank. At this time, two nonnatal males Y and X, with little antagonism from Shan, gradually began to work their way into the troop. Y slowly gained the support of middle- to lower-ranking females and the young males (Hazama 1972), and Shan left the troop in August 1961, apparently after an increasing lack of support from troop members (Hazama 1972). Around No- vember of that year Y became alpha. He was the first known nonnatal male to become alpha in the Arashiyama group. He remained alpha male until leaving the group four years later in September 1965 (Hazama 1972; Koyama 1970). In November 1962, the alpha female Tokiwa died. Matsu, thought to be Tokiwa’s daughter, became the next alpha female (Hazama 1972). As of March of that year, the troop had grown to a size of 102 individuals (Norikoshi and Koyama 1975). Consorts, Kin-Group Rank Change, and Troop Fission In March of 1966 the troop had reached 163 members and in June it fis- sioned. This was the first case of fission observed in a Japanese macaque troop for which the genealogical relations were known (see Koyama 1967,28 Huffman FIGURE 3. CHRONOLOGY OF GROUP FISSION. as ae? ? ; 1948 MARS. 1955 - ZAO i Tokiwa - LINCOLN 1960 SHAN - Y - Matsu 1965 - ZOLA gm Mino AO pyMatsu - w 1970 Ma-59 - Rn-582 1975 Mi-632 - Mi-63692 1980 Mi-6369742 - K-63 1985 - D-648 pyMi-6369743% Mi-73% mGi69g - [ Joreerer’ 1989 Arashiyama Chronology of group fission and the transfer of alpha male and fe- male positions of the Arashiyama A, B, E and F groups since 1948. C and D groups are excluded, but see text for description. 1970; Norikoshi and Koyama 1975 for a detailed description). According to Koyama (1970), fission was the result of a breakdown in the normally sta- ble kin-group rank order. He traced the beginnings of this breakdown to September 1964 when X, the beta male, disappeared and Ao (son of the middle-ranking Betta) entered the central part of the troop as a subleader. During this period (before the alpha male Y's disappearance), agonistic en- counters were observed among the higher-ranking individuals in the centerHistory of the Arashiyama Japanese Macaques 29 of the troop. Notable were the attacks on Zola and Ao by the younger Ma- 598 backed by his mother Matsu and sisters. According to Koyama (1970), Zola had been the third-ranking male under Y and X. After X then Y disappeared, Ao temporarily took leader- ship of the troop. Koyama notes that shortly thereafter Zola became alpha male and with his help Ao was able to dominate Ma-59d and Matsu. Strong social bonds developed from consorts between Ao and Matsu. This led to instability in rank relations and a resultant loss of unity among kin- groups. Mino, then a mid-ranking female, began associating with Zola. With rank dependence on Mino and Zola, lower-ranking females rose in rank above females of higher-ranking kin-groups. Similarly, Mino, through her rank dependence on Zola, became head of her kin-group. This came about after a series of attacks on her aged mother Kojiwa (Koyama 1970). From within the same kin-group, the status of alpha female was transferred from mother to daughter three more times, twice (i.e., Mino to Mi-63?, Mi-632 to Mi-6369@) as the result of aggressive conflict. In the case of Mi-632 to Mi-63692, Mi-639 died as an indirect cause of injuries re- ceived from an attack by her daughter and her granddaughter Mi-636974 9 the same day (N. Asaba 1984, park records). The following year, Mi- 63692 was removed by the park management because of her highly aggres- sive behavior toward people. Her daughter Mi-636974 then became alpha female (fig. 3). Reorganization of kin-groups into A and B troops The outcome of this fis- sion, which lasted until October of 1966, was the formation of two stable and independent groups; the dominant B troop led by Zola and Mino, and the A troop led by Ao and Matsu as alpha males and females (fig. 3). The females of the middle- and lower-ranking kin-groups below and inclusive of Mino’s moved into B. The females of the originally middle- and higher- ranking kin-groups above Mino moved into A. Almost all males over 5.5 years of age moved into the troop in which their female relatives did not reside. Thus, Ma-59¢ left his mother, the alpha female, in A troop and joined Zola as beta male of B troop (Koyama 1970). Ao left A troop in July of 1967, after 18 months as alpha male. He was replaced by W. Dai, a close associate of W’s in the periphery of the group since 1964, then became the beta male (Norikoshi 1977). W remained alpha male of A troop up through February of 1972, when the entire group was captured and sent to Texas. (Details of capture and transfer follow, and also see Fedigan, this volume, for the history of the A troop in Texas.) Accord- ing to Mano (unpublished data), Matsu gradually weakened and began to spend most of her time in the periphery with her daughter Ma-64°. By March 1971, the Rheus kin-group was dominant over the Matsu kin- groupand Rh-58 was the alpha female. It is not known whether or not30 Huffman Rh-58? was dependent on W for her rank at this time, but in 1972 after the group went to Texas, she appeared to maintain her rank independently of Dai (Mano, unpublished data) who had become alpha male on the first day of their release (Fedigan, this volume). Zola left in July of 1970 and Ma-59d became the next alpha male of B troop (Norikoshi 1977), remaining so for a record 11 years. Changes in Kin-Group Rank Order, Female Subgrouping, and Fission From November 1977, after a second feeding site was set up, subgrouping by several mid- and low-ranking females was observed. These females of the Kusha, Rakushi, and Ai kin-groups began to use the site after being outranked by Glance and her daughters (see Takahata 1982a). By February 1978, 12 members of the Kusha lineage (four adult females and their off- spring) had formed a stable group with two adult peripheral males Koj- 6269¢ and GI-6073¢. This independently moving unit was named C group (Grewal 1980a; Takahata 1982a). The group was captured between March 1978 and January 1979 (excluding two three-year-old Kusha males who es- caped) and sent to the Choju City Zoo, South Korea (Asaba 1984, park records). As of March 1978, B group had reached 258 individuals (Koyama et al. 1980). Feeding at these two sites was continued and in May of 1978 the park staff recognized a second independently traveling group. The park later named the resultant, approximately 28-member group, D troop (Suzuki, Matsuura, and Kawamichi 1987). According to Asaba (personal communi- cation) six mid- to low-ranking females and their offspring of the Rakushi, Ai, Glance, and Blanche kin-groups traveled with adult males, Solitary 76- 13 (confirmed by Dr. Atsuya Yoshida, of Osaka University, to have been a nonnatal male of Minoo troop, Osaka), Koj-6273d, and Mo-6773¢ in the periphery. All but Ai-6166?, -759, -762, and -78d were captured in late July. From this group, including Solitary 76-1d, 15 individuals were se- lected and sent to the Kaibara Family Land, Hyogo. Ai-61662 and her offspring were captured in early August and successfully reintroduced back into B group (N. Asaba, personal communication). A second artificially formed group of 23 individuals, captured at Arashiyama in 1981 (Arash- iyama PRI group) and housed in an outdoor enclosure at the Kyoto Univer- sity Primate Research Institute, has sometimes mistakenly been referred to as the Arashiyama D group in the literature. Alpha Male Instability, a Lack of Female Support? K-63d became the next alpha male after Ma-59¢ left the group in June 1981 (park records). While adult male rank relations remained stable, K-63¢’s relationship with theHistory of the Arashiyama Japanese Macaques 31 alpha female was often ambiguous (Huffman, unpublished data). Part of the cause of this ambiguity was the alpha female’s aggressive attitude toward his relentless following of her. K-63 0's unstable position was obvious from the considerable time and energy he spent trying to consolidate sexual and nonsexual relations with her between 1983 and 1985. In general, female support for him was at an all-time low as evidenced by their unwillingness to form consort relationships with him (Huffman, this volume). K-63¢ left the troop five times between 1981 and 1986 during his ten- ure as alpha male, for periods of ten days to two months (park records). The lack of female support seems to have had an influence on K-630's instability in the troop. A similar trend, although not as drastic, was ob- served for Ma-59d¢ during the 1979-80 mating season (Huffman 1981), little over a year before he left the troop. Both males were recognized to have had high frequencies of mounting series in previous years (Stephenson 1975; Takahata 1982a, 1982b; Huffman, unpublished data). _ Growing Female Support, Coexistence with Female Relatives, and Troop Fission In January of 1986 the approximately 246-member troop again began to show signs of fissioning (H. Suzuki, unpublished park records). A detailed account of this fission and the events leading up to it are to be prepared in the future by Suzuki. However, due to the importance of this event to the group’s social history, the following brief accounts extracted from park records and my own observations follow. According to N. Asaba (unpublished data), signs of disunity first ap- peared after the death of the beta male Ran-63d in December of 1985. Also of importance to this fission, was the growing social influence of two males Mi-736 and Co-6573d (fourth- and fifth-ranking males). These males had reached social maturity and were becoming influential figures within the troop social network. An indication of this was the high fre- quency of mating activity and the number of multiday consorts for both males during the 1984-85 mating season (Huffman, this volume). Mi-73 had begun to attain female followers, such as GI-69?, from the fall of 1984. Furthermore, these two males’ relatives made up two of the highest- ranking kin-groups in the troop. Given the examples of the fission into A, B, and C groups in which aggressive conflicts between an adult male and females of the same kin- group were observed and in which they moved into different troops (Koyama 1970; Norikoshi and Koyama 1975; Grewal 1980a), the coexist- ence of Mi-736 and Co-65736 with their relatives could be considered a cause of future social friction and instability. Had these males not socially32 Huffman integrated so well with certain adult females, perhaps they would have left the group on their own. However, this was not the case and the group began to grow apart. Adult Male Group Identity and Subgrouping The division of male leadership into two groups became apparent with the sudden rise of the peripheral male Op-6076¢, from sixth to fourth rank, above Mi-73d and Co-65733. Op-6076d was beginning to spend more time in or near the center of the troop antagonizing adult females (park records; Huffman, un- published data). It can be argued that Op-6076d was identifying himself with the dominant group of K-63d, by responding to Mi-73d and Co- 65736 as if they were members of a different, subordinate group. Mi-73¢ and Co-6573¢ responded subordinately to him and were spending less time near the group (park records). Given the disparity in age, social experience, and body size between Op-6076¢ and these two older males, the best ex- planation for their reversal in rank appears to be that the group was under- going a reorganization of membership into two new groups, K-63¢’s group being dominant to the newly forming group of the allied males Mi-73d and Co-65736, and their female followers. On the morning of May 29, according to N. Asaba (personal commu- nication), the subgroup of Mi-736 and Co-6573¢, and their growing num- ber of followers were chased out of the feeding station by individuals of the main group. Two separate feeding sites were established on June 12, to insure that both groups had access to the provisioned food so that Mi-73d’s group would be prevented from moving down into the edge of the city. On August 16, N. Asaba confirmed that the two groups were sleeping in dif- ferent sites at night. After consulting with N. Koyama, the two groups were officially named E and F troop (fig. 3). K-63d and Mi-636974? were the alpha male and female of E troop. Mi-73d became alpha male of F troop, and GI-69? through her rank dependence on him as a follower since 1984, became alpha female. Reorganization of Kin-Groups into E and F Troops According to Suzuki et al. (1987), as of September 1, 1986, E group consisted of 149 individuals (73 females, 76 males) including all the females of the Kojiwa, Mino, Coo- per, Yun, Chonpe, and Kusha kin-groups, most of the Blanche group fe- males, and 1] females from the lower-ranking Oppress, Rakushi, Shiro, Momo, Glance, Shirayuki, and Meme kin groups. F group consisted of 97 individuals (58 females, 39 males) including almost all of the females of the Oppress, Shiro, Momo, and Glance kin-groups, plus the females, Ra- 586675? and BI-5870@. All males 2.5 years or younger, excluding or- phans, remained with their mothers. Although at this stage male membership was not yet stable, most males 4.5 years or older were moving out of the group in which their female relatives resided (Suzuki et al. 1987).History of the Arashiyama Japanese Macaques 33 On October 4, K-63 and beta male De-64¢ left the group for two months, returning together on December 1 (park records). During their ab- sence, the alpha female Mi-636974? was the highest-ranking individual in E group. Upon their return, K-63’s and De-646’s previous rank positions were not challenged by other males. Toward the end of December, 1986 K-63d was outranked by De-64¢ after sustaining dehabilitating injuries from a hunter’s snare. De-64d and Mi-636974¢ have remained the alpha male and female through to the present (1989). K-63d next left the group in June 1987 for six months, to return again in December. According to R. Asaba, K-63d was not able to enter the center of the group and was outranked by young juvenile males in the periphery. He left for good on December 23, 1987 (park records). In the 1987-88 mating season, due to a rise in rank precipitated by a long consort with Mi-73d, Op-607580 rose in rank above GI-69? to be- come the new alpha female. Op-607580? has maintained this status to the present (1989). POPULATION DYNAMICS In February 1964, 22 monkeys (16 females, six males) were temporarily captured and examined, and tests for tuberculosis, dysentery bacillus, Sal- monella bacillus, and B virus were negative. With the exception of intesti- nal parasites in 17 individuals and one case of pyorrhea in a ten-year-old female, the group was in good health (Tanaka 1964). With the exception of one female, Mino, who gave birth at the age of 25, most females are reproductively active between the ages of four and 23. According to Koyama et al. (in prep.) for females between the ages of five and 19, the reproductive rate ranges from 46.7% to 69.0%, respectively. By 23, this rate decreases to 11.1% (n = 827 females of known age). Between 1954 and 1983 a total of 975 infants were born into the Arashiyama troop (1954-65), A and B group (1966-72), and B group (1972-83). Calculating from this (excluding 20 stillbirths of unidentified sex) the male (n = 486)- female (n = 469) sex ratio at birth is 1:1.04. Interbirth intervals were cal- culated from 770 samples of 172 females who gave birth to two or more infants. An interval of one year is most common (57.5%, n = 443) fol- lowed in succession by intervals of two years. (38.6%, n = 297), three years (3.6%, n = 28), four years (0.1%, n = 1), and six years (0.1%, n=1). During the 18 years between 1954 and 1972 the group (A and B com- bined) increased in size from an estimated 28 to 301 individuals. This amounts to a 13.7% increase per annum (Koyama 1984). Constraining this growth are several factors such as death and male transfer. Due to the fact34 Huffman that males emigrate at an early age from their natal group and others im- migrate as adults, it is impossible to accurately estimate their life span. For females, however, this is not the case and at Arashiyama the oldest re- corded female (as of July 1989), Mino, is 32 years old. This is exception- ally old and only two other females (Yun and Oppress) have been known to reach 30 years of age. Between 1954 and 1984 a total of 203 deaths were recorded, 46.3% of which were represented by infants less than one year of age. During this same period, 146 males were believed to have emigrated and an additional 432 males and females were captured (Koyama 1984). In contrast to the large number of emigrations, permanent immigration is extremely low (n = 14) accounting for only 1.4% of the total population increase between 1954 and 1984 (Koyama 1984). During the mating season a few unknown males visit and are observed to mate with group females. However, like most other groups, Arashiyama is isolated from its nearest neighbor by several kilometers due to the loss of habitat, i.e., the creation of no-monkey zones by commercial forestry and the expansion of cities and agricultural areas throughout Japan. This makes male transfer difficult. In spite of this obstacle, several cases have been observed in which males from Minoo group have been found in the Arashiyama group, nearly 30 kilometers away, and vice versa (park records). It is interesting to note that the longest recorded distance traveled by a male Japanese macaque in Japan is 60 kilometers from his natal group (Fukuda, Takanaka, and Muramatsu 1974). TRANSPLANTING ARASHIYAMA A GROUP TO TEXAS In 1968 two American researchers, Steven Green from Rockefeller Univer- sity and Gordon R. Stephenson from the University of Wisconsin, came to Arashiyama to carry out research on vocal communication and the social structure of mating behavior at Arashiyama (Green 1975a, 1975b, 1977; Stephenson 1973a, 1973b, 1975). They came to Japan under a Japanese- American cooperative research project directed by John T. Emlen of the University of Wisconsin and Ryoji Motoyoshi of Kyoto University. On the Japanese side, Munemi Kawabe and Kawamura of the Osaka City Univer- sity assisted in the research and management of research-related business (Koyama and Norikoshi 1984). A by-product of this cooperative project were the plans made to send A group to the United States, and to continue comparative studies between these two genetically similar groups in isolation from each other. According to Koyama and Norikoshi, the process of sending A group to the United States was a difficult, expensive, and time-consuming project. In December of 1968 a group of ten American scientists; J. T. Emlen, C. R. Carpenter,History of the Arashiyama Japanese Macaques 35 C. B. Koford, J. King, E. Banks, E. Sackett, W. Mason, P. Marler, C. Southwick, J. Vandenburg, and B. Alexander, was formed to investigate the logistics of such a project. The first meeting was held at the Argonne Na- tional Laboratories in Illinois. Several conditions were made as fundamental requirements for a place to relocate the troop. They were that (1) the terrain should be hilly and forested, (2) the climate should be moderate, (3) the area should be en- closed so that the monkeys cannot escape, (4) the site should be easily accessible from more than one major university center, and (5) there should be no outside disturbance to the troop. Four suggested locations that met these requirements, Argonne National Laboratories, Fort Custer National Forest in southern Michigan, an offshore island along the coast of Georgia or South Carolina, and an isolated forest in Hawaii, were investigated. However, these sites were eventually rejected by the group. Next a site in Columbia, Missouri, was suggested and plans to build a fence and receive the monkeys by January of 1971 was tentatively set. However, funding to build the fence did not come through. The date was extended to July of 1971 and in May of that year the American side applied for funds from the National Science Foundation (Koyama and Norikoshi 1984). Back in Japan, the subordinate A group occasionally moved down to the edge of the town, causing problems for the local people living along the edge of the mountains. Complaints from these people increased and so Iwata planned to construct a fence around A group’s feeding site in order to capture and hold them until a site was established in the United States, and the group could be sent over. However, capturing was delayed and yet an- other site was found and had to be abandoned at the last moment. The strenuous efforts of both sides to complete the project continued (Koyama and Norikoshi 1984). Two researchers, Tetsuzo Mano, a student of Osaka City University and Tim Clark, a graduate student working under Emlen, were selected to con- duct preliminary observations of A group in Japan and to accompany the group to the United States (Mano, Norikoshi, and Koyama 1972; Clark and Mano 1975). Clark came to Arashiyama in December of 1971 (Koyama and Norikoshi 1984). The future of A group seemed uncertain, as a suitable site had not yet been found. However, the problem was soon to be solved through the ef- forts of Claud Bramblett at the University of Texas in Austin. An agreement was made with Edward Dryden Jr., a rancher in Laredo, Texas, to receive the group and to provide the necessary land and fencing required to provi- sion them, on the condition that in the future he would use the expanding excess population as the core for a commercial reproductive colony (Koyama and Norikoshi 1984).36 Huffman Now that the United States site was finally decided, capturing began on February 16, 1972. On the 16th, 125 individuals were caught on the first attempt. Several attempts were made and capturing continued until the 20th, when 151 of the 158 members of A troop, plus a solitary male, were finally captured (Koyama and Norikoshi 1984). Under the direction of Mitsuo Iwamoto of Kyoto University’s Primate Research Institute, 14 scientists examined each individual, recorded body weights, and were able to give all but one a tattoo for future identification. One individual died and a hybrid Rhesus x Japanese monkey (Rhotte- 61689) was removed, making a total of 150 individuals; 94 females, 56 males (Mano, Norikoshi, and Koyama 1972). On February 22, 172 the Arashiyama A group was sent off from Osaka to their new home in Laredo, Texas, where Clark and Mano (1975) observed the group’s adaptation to the new environment. The researchers in Texas presently refer to A group as ‘*Arashiyama West’’ in the literature (see Fedigan, this volume, for details on A group’s adaptation to the Texas site). RESEARCH AT ARASHIYAMA Certainly the most valuable aspect of the Arashiyama macaques is the long- term data composing the genealogical records and individual life histories accumulated by the cooperative efforts of over four generations of research- ers and park staff since 1954. Among the numerous other monkey parks opened in the 1950s, comparable data are available only at Koshima and Katsuyama. Table 1 gives the names of the researchers who worked and published on the Arashiyama macaques. Being a free-ranging population, studies on the group largely have been restricted to naturalistic observations. This research can be categorized into six major areas including social dynamics and kinship, sexual behavior, population dynamics, feeding ecology, vocal communication, and behav- ioral development. Specific topics investigated cover a variety of areas ranging from troop fission, male transfer (Koyama 1970; Norikoshi and Koyama 1975), reproductive biology, rank and reproductive success (Taka- hata 1980, 1982b; Huffman 1987), paternal care (Wolfe 1981a; Grewal 1984), and the transmission of cultural behavior (Green 1975a; Huffman 1984a, 1986a, 1986b). Of the small amount of research done on temporarily captured mon- keys, most notable was the multidisciplinary study done in February 1964 (Tanaka 1964). The results attained from this project were used as compar- ative data for similar work done at other sites in the areas of morphology, hematology, and cardiology (see Nigi, Tanaka, and Noguchi 1967; Tanaka and Nigi 1967; Iwamoto 1967, 1971; Noguchi et al. 1969).History of the Arashiyama Japanese Macaques 37 TABLE 1. CHRONOLOGY OF RESEARCHERS AT ARASHIYAMA Researcher Period of Fieldwork Current Affiliation Itani, J. 1951 Retired Kawamura, S. 1951 Retired Hazama, N. 1951 Deceased July 1954-Nov. 1961 Nakajima, K. 1955-63 Japan Birds of Prey Research Ctr, Hirosei, S. 1955-58 Japan Monkey Ctr, Koyama, N. Mar. 1964—Mar. 1967 Kyoto Univ. Feb. 1972-periodically to present Norikoshi, K. Jan. 1967-Feb. 1972 Jochi Univ. Mano, T. Apr. 1969-Feb. 1972 Torigoe Jr. High School Green, S, July 1968—-Aug. 1969 Univ. of Miami Stephenson, G. July 1969-Oct. 1969 Univ. of Wisconsin Nishida, K. 1973-1974 Kyoto Univ. Koyama, T. Spring 1974 Ochanomizu Univ. Grewal, B. S. Oct. 1975-July 1978 Guru Nanak Dev Univ. Takaragawa, N. May 1975-Mar. 1976 Chiba City Zoo Takahata, T. Oct. 1975-Sept. 1977 Kyoto Univ. Wolfe, L. Aug. 1977-July 1978 Univ. of Florida Huffman, M. A. July 1979-Sept. 1980 Kyoto Univ. Oct. 1983-Mar. 1985 Sept. 1986—periodically to present Suzuki, H. & May 1981-Dec. 1981 Iwatayama Monkey Park Matsuura, N. _ Suzuki, H. Jan. 1983-1989 Full-time employee Iwatayama Monkey Park Inoue, M. May 1981-Sept. 1985 Himeji Gakuin College Nakamichi, M. Apr. 1983-July 1983 Osaka Univ. Quiatt, D. Jan. 1984 Univ. of Colorado Note: Revised and updated from Baldwin, Koyama, and Teleki 1980. Contributions of Research at Arashiyama to Primatology Several of the studies done at Arashiyama have had a particular and lasting influence on the direction taken by primatology (table 1, references, and bibliography, this volume, for full details). Perhaps Hazama’s greatest contribution to primatology was his timely observations of male transfer from Arashiyama to Hieizan (Hazama 1965) and the emigration of nonnatal males into the Arashiyama troop (Hazama 1972). As mentioned, at that time, Japanese macaque society was believed38 Huffman to be closed. In 1958, Hazama started his study of the Hieizan group about 18 km away on the eastern side of Kyoto (fig. 1), periodically returning to Arashiyama (Hazama 1964). During this period he noted the disappearance of subadult male, Chikusha, son of the alpha female Tokiwa. Five months later on January 19, 1960, Hazama found this male in the Hieizan group (Hazama 1965). This was the first confirmation of a male transfer from his natal group to another in Japanese macaques. Supportive observations were made at Shigakogen (Wada 1964; Yoshihiro and Tokida 1976). Nishida (1966), based on his own observations at Takagoyama and on an overview of the available data from other groups, concluded that for males, leaving their natal group is an important phase in their life history. From these observations and others it became obvious that Japanese macaque groups are perpetuated through matrilineal groups (Itani 1972, 1977, 1985b; Sugi- yama 1976). The next major study at Arashiyama was done by Naoki Koyama, a graduate student at Osaka City University and a student of Kawamura’s. Based on the detailed records of intraindividual kin relationships left by Hazama and Nakajima, Koyama conducted detailed studies of dominance rank order (Koyama 1967, 1970) and playmate relations (Koyama 1985). His work provided the first detailed analysis on kinship dynamics in rela- tion to group fission; elucidating the correlations between kinship, social relationships, and behavior, which have greatly influenced the direction of current primatology. Norikoshi, also a student of Kawamura’s, joined Koyama in January 1967 and together they investigated the dynamics of group fission (Koyama 1970; Norikoshi and Koyama 1975). In this classic study they followed the complex interactions between males and the matrilineally based female groups during fission. In doing so they elucidated the matrilineally based group infrastructure of Japanese macaques and provided a model for incest avoidance and a mechanism for intergroup gene flow. Between 1967 and 1972, the coexistence of these two daughter groups at Arashiyama provided the opportunity to study male transfer based upon a complete knowledge of kinship and dominance rank. In particular, Noriko- shi made significant contributions to our knowledge of the development of mother-son and peer-mate relationships and their effects on male solitariza- tion (Norikoshi 1974a, 1974b, 1974c, 1975a, 1975b, 1977). Ttani (1983) characterized the mid 1970s as the beginning of a phase in primatology in which several important problems proposed by previous re- search were solved. One of the problems addressed was incest avoidance. Several explanations had been proposed, including mother’s dominance over her son, male transfer, and psychological mechanisms (eg., Itani and Tokuda 1958; Sade 1968; Kawai 1969; Itani 1972; Norikoshi 1974a, 1974b, 1974c).History of the Arashiyama Japanese Macaques 39 At Arashiyama, Baldev S. Grewal and Yukio Takahata, both graduate students at Kyoto University, further elucidated a psychological mechanism for incest avoidance. Grewal (1980a) demonstrated that adult males remain- ing in the group with female kin relatives tend not to associate with them. Takahata (1982b) then demonstrated that the avoidance of copulation was Statistically significant between individuals up to within the third degree of consanguinity. The existence of close persistent relationships in certain male-female pairs were first described by Kitamura (1977) at Takasakiyama. At Arash- iyama, Takahata (1982a) developed this further and found that mating rela- tions played an important role in the formation of such intimate relationships, which he called ‘‘peculiar proximate relations’’ (Takahata 1982a). He found that such pairs tended to avoid mating as if they were kin-related and suggested that affinity (psychological bonding) may act to suppress the sexual drive between them (Takahata 1982a, 1982b). Itani (1985b) suggested that the mechanism of incest avoidance as made clear at Arashiyama, was deeply involved in the formation of social structure. The most influential aspect of the research done at Arashiyama is the complete long-term genealogical records. This has made it possible to con- duct detailed studies from a sociological and demographic perspective not currently possible for many other primate populations. Perhaps the unique- ness of Japanese primatology is in part reflected by this diachronic ap- proach. Itani (1985b) suggested that primatologists should not only observe behavior from a synchronic standpoint (i.e., the study of mechanisms of behavior at a given time), but also from a diachronic one, by studying the life history of individuals and then ultimately the sociological history of the troop in terms of change over time (also see Asquith, this volume). Development and Management of the Iwatayama Monkey Park The Iwatayama Monkey Park was officially opened to the public on March 20, 1957, three years after the first plans for its construction were made. In October of that year, a group of rhesus monkeys was put in an open enclo- sure part way up the mountain to help compensate for the unpredictable appearances of the Arashiyama group to the feeding station (Hazama 1962; Asaba 1984). This group of rhesus monkeys was maintained until around 1969 (Koyama personal communication). During their presence at the park, two cases were observed in which a Japanese macaque female climbed into the enclosure, mated, and produced viable offspring (Mano, Norikoshi, and Koyama 1972; Wolfe 1981b). As the park was a tourist spot, a concession stand was opened and people were allowed to feed the monkeys peanuts.40 Huffman The orientation of the park as a tourist area under Iwata’s management continued with little change until 1974. Financial Difficulty and a Change in Ownership By the early 1970s the park was facing financial difficulty, as were many other monkey parks. In April of 1974, Kyoto city purchased the land from Iwata, declaring it a protected historical reserve, but the financial responsibility of management was left to Iwata. There was a feeling of uncertainty about the future of the monkeys. On March 4, 1974, a month before the land was officially bought, a meeting of the Arashiyama Natural History Research Group was held. They made a proposal to Kyoto city officials to turn Iwatayama into a nature observatory and offered Iwata their assistance. Kawamura, Koyama, and Norikoshi had previously written an article in the group’s first report (1973) in which they expressed the importance of continuing research at Arash- iyama and discussed the duty of researchers not only to work toward the advancement of science, but also to put their findings into the reach of the public. Kinji Imanishi expressed support for further research at the park and extended the responsibility of protecting and supporting the park to the citizens of Kyoto. The scientists’ actions had a significant effect on the future development of the park. In February 1976, Nobuo Asaba and his family took over the manage- ment. He started by making plans for its renovation. First the 600-meter trail climbing 125 meters up the side of the mountain was widened so that a small motor transport could carry provisions for the monkeys, concession supplies, and building materials to the feeding station (fig. 2). In order to operate the park and provide provisions for the troop, an entrance fee was charged, as before, and the concession stand for tourists was continued. However, visitors were no longer allowed to feed the monkeys out-of-doors; fruit and vegetables provided by the park could be given to monkeys only from inside the visitors’ room through wire netting. Development into a Scientific and Educational Resource In the spring of 1979, Asaba constructed a two-room building (Arashiyama Natural History Research Station and Museum) near the feeding site to serve as a gathering place and to house a nature exhibit for the public. A small library was assembled with a collection of literature on primatology, including all the reports and papers of research done at Arashiyama and field guides to the flora and fauna of the area. From that point, the orientation of the park took a new direction re- flecting the efforts of the Arashiyama Research Group and Asaba's con- cern for maintaining and preserving the scientific value of the Arashiyama macaques. Up to this point, the knowledge of individual identification of the monkeys was passed down from one researcher to another. Shortly after Asaba took over management of the park, the staff took responsibility forHistory of the Arashiyama Japanese Macaques 41 maintaining the troop records and genealogies. With help from Takahata, Asaba and his assistants (some of whom were trained primatologists work- ing on a temporary basis) learned the names of all the individuals. The use of simple tattoos on the face and ears, first begun at Arashiyama in 1964, was used in order to insure individual identification, in particular of young males who spend most of their time in the periphery of the group or leave it completely at an early age. Tattooing made it easier for new researchers to recognize the animals, and proved invaluable in the identification of males who had left the group and had appeared elsewhere. Also, with the assis- tance of the staff in identifying all individuals of the group, it became fea- sible for researchers of other groups to come and do short-term comparative studies (e.g., T. Koyama, Fujii, and Yonekawa 1981; Nakamichi 1984a, 1984b). Now every year researchers from Kyoto University, Osaka City University, and Doshisha University, conduct undergraduate seminar field courses at Iwatayama. The students are instructed in the basics of ethology and animal observation. This transformation allowed for a wider use of the group as an educa- tional resource. One of the main objectives was to educate the public about the monkeys and their habitat and to instill an appreciation of the environ- ment and the need for its conservation. The Arashiyama Nature Club was established by the park in May of 1979 to help fulfill these objectives. The club provides opportunities to the membership such as an annual summer workshop, focusing on some aspect of primate behavior that helps them to learn how to identify the monkeys. As of April 1987 the club had 162 of- ficial members. A bulletin, the Hoobukuro (‘‘check pouch’’) is printed ev- ery other month and includes information about group activities such as bird watching, nature hikes, essays, and drawings by club members, and a chronological reference of happenings within the troop as observed by the staff, club members, and researchers. These activities have helped to make the neighboring people, most of whom only identified the monkeys as pests, more aware and sympathetic to their presence. Present Conditions and Some Changes Since Provisioning Provisioning ac- celerates population growth (Sugiyama and Ohsawa 1982), at least in the short term (Asquith 1989). In an attempt to curb this growth, several mon- key parks throughout Japan, including Arashiyama, experimented by de- creasing the amount of provisioned food. At Arashiyama, however, this led to increased problems with hungry monkeys moving into the edge of the city during the daytime and causing problems with local shops, houses, and hotels (Asaba 1984). Although not a preferred practice of the park, captur- ing monkeys has been practised to help alleviate overpopulation. Provisioning has undoubtedly affected the group’s feeding habits. Since the troop shifted permanently to the Iwatayama area after 1955, its home range has decreased from eight square kilometers to an estimated .7 square42 Huffman kilometers (Hazama 1972; Asaba 1984). This concentrated use of space by the large population resulted in severe foraging pressure on the vegetation around the feeding station. During a three-year survey started in 1954, Hazama recorded 192 plant species eaten by the monkeys throughout their home range (Murata and Hazama 1968). Twenty-two years later, a preliminary 12-month survey of B group cataloged 67 plants from 34 families, eaten in the vicinity of the feeding station (Huffman 1984b). Interestingly, this list included 17 species not observed by Hazama, suggesting that the group may have acquired at least some of these new food items after provisioning. Direct comparison between these two surveys is not possible due to the preliminary nature and restricted area of observation of the latter. However, it can be said that the macaques are still partially reliant on the natural vegetation. At a time when loss of habitat and persecution as pests threatens the remaining isolated populations of Japanese macaques, steps to restore some of their habitat and to develop alternative methods of crop-raiding control are urgently needed. There may never be another chance to repeat what has been done at Arashiyama. Hopefully this paper has in part preserved the history of this invaluable treasure, and will in some way contribute to its future protection and development. ACKNOWLEDGMENTS I wish to give my sincerest thanks to Nobuo Asaba, his two sons Ryosuke and Shinsuke, Ms. Hisayo Suzuki, and Ms. Kazubo Kanbara of the Iwatayama Monkey Park for their generosity and help in my studies, in providing access to park records, and for their endless patience in answer- ing my endless questions. Special thanks must also go to Drs. Junichiro Ttani, Naoki Koyama, Toshisada Nishida, and Yukio Takahata and to Mr. Tetsuro Mano for their support, criticism, and advice. This paper is dedi- cated to all the monkeys of Arashiyama, past and present, and to those people without whose efforts this history would never have been recorded and who have contributed so much to our understanding of the subtleties of primate social life.Stone-play began among ju- veniles in 1979, and is now common at Arashiyama, where it has been the subject of study by the author. Cour- tesy of Michael A. Huffman The provisioning grounds at Arashiyama, Japan, with a view of Ky- oto in the background. Courtesy of Michael A. HuffmanThe Arashiyama provisioning grounds in winter with monkey in the foreground, Kyoto in the background, and Mount Hiei in the dis- tance. Courtesy of Michael A. Huffman Evening feeding grounds, Arashiyama, Kyoto. Courtesy of Pamela AsquithHistory of the Arashiyama Japanese Macaques 45 REFERENCES Asaba, N. 1984. 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