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The Japanese macaque ( Macaca fuscata), also known as the snow monkey, is a terrestrial Old World monkey species that is native to . Colloquially, they are referred to as "snow monkeys" because some live in areas where snow covers the ground for months each year – no other non-human lives farther north, nor in a colder climate. Jigokudani Monkey Park, Nagano: Explore the Heart of Japan Individuals have brownish grey fur, pinkish-red faces, and short tails. Two subspecies are known.

In Japan, the species is known as Nihonzaru (ニホンザル, a combination of Nihon 日本 "Japan" + saru 猿 "monkey") to distinguish it from other primates, but the Japanese macaque is very familiar in Japan—as it is the only species of monkey in Japan—so when Japanese people simply say saru, they usually have the Japanese macaque in mind.


Physical characteristics
The Japanese macaque is sexually dimorphic. Males weigh on average , while females average .Fooden J, Aimi M. (2005) "Systematic review of Japanese macaques, Macaca fuscata (Gray, 1870) ". Fieldiana: Zoology 104:1-200. Macaques from colder areas tend to weigh more than ones from warmer areas.Hamada Y, Watanabe T, Iwamoto M. (1996) "Morphological variations among local populations of Japanese macaque ( Macaca fuscata) ". In: Shotake T, Wada K, editors. Variations in the Asian macaques. Tokyo: Tokai Univ Pr. pp. 97–115. The average height for males is , while the average female height is . The weight of their brain is approximately . Japanese macaques have short stumps for tails that average in males and in females. The macaque has a pinkish face and posterior.Rowe N. (1996) The pictorial guide to the living primates. East Hampton (NY): Pogonias Pr. pp. 124–5. The rest of its body is covered in brown or greyish hair. The coat of the macaque is well-adapted to the cold and its thickness increases as temperatures decrease. The macaque can cope with temperatures as low as −20 °C (−4 °F).

Macaques mostly move on all fours. They are semiterrestrial, with females spending more time in the trees and males spending more time on the ground. Macaques are known to leap. They are very good swimmers and have been reported to swim a distance of more than half a kilometer. The lifespan of Japanese macaques is up to 32 years for females and up to 28 years for males, which is high when compared to what typically is seen in other species.


Behavior

Group structure
Japanese macaques live in societies, and females stay in their natal groups for life, while males move out before they are sexually mature. Macaque groups tend to contain adults of both sexes. In addition, a Japanese macaque troop contains several matrilines. These matrilines may exist in a dominance hierarchy with all members of a specific group ranking over members of a lower-ranking group. Temporary all-male groups also exist, composed of those who have recently left their natal groups and are about to transfer to another group. However, many males spend ample time away from any group,
(1976). 9780120045075
and may leave and join several groups.

Females of the troop exist in a stable dominance hierarchy and a female's rank depends on that of her mother. Younger females tend to rank higher than their older siblings.Takahata Y. "Diachronic changes in the dominance relations of adult female Japanese monkeys of the Arashiyama B group". In: Fedigan LM& Asquith PJ, editors. The monkeys of Arashiyama: Thirty-five years of research in Japan and the west. Albany(NY): SUNY Pr. p123-39. Higher-ranking matrilines have greater social cohesion.Koyama NF. (2003) "Matrilineal cohesion and social networks in Macaca fuscata". Int J Primatol 24(4):797-811. Strong relationships with dominant females can allow dominant males to retain their rank when they otherwise would not. Males within a group normally have a dominance hierarchy, with one male having alpha status. The dominance status of male macaques usually changes when a former alpha male leaves or dies.Sprague DS, Suzuki S, Tsukahara T. (1996) "Variation in social mechanisms by which males attained the alpha rank among Japanese macaques". In: Fa JE, Lindburg DG, editors. Evolution and ecology of macaque societies. Cambridge (UK): Cambridge U Pr. p 444–58. Other ways in which status of male hierarchy changes, is when an alpha male loses his rank or when a troop splits, leaving a new alpha male position open. The longer a male is in a troop, the higher his status is likely to be.

Females typically maintain both social relationships and hygiene through grooming. Grooming occurs regardless of climate or season. Females who are matrilineally related groom each other more often than unrelated individuals.Koyama N. (1991) "Grooming relationships in the Arashiyama group of Japanese monkeys". In: Fedigan LM, Asquith PJ, editors. The monkeys of Arashiyama: thirty-five years of research in Japan and the west. Albany (NY): SUNY Pr. p211-26. Females will groom unrelated females to maintain group cohesion and social relationships between different kinships in a troop. Nevertheless, a female will only groom a limited number of other females, even if the group expands. Females will groom males, usually for hygienic purposes, but that behavior also may serve to attract dominant males to the group. Mothers pass their grooming techniques to their offspring, most probably through social rather than genetic means, as a cultural characteristic.


Documented female troop leadership
Yakei is a female who rose to leadership of her troop at Takasakiyama Natural Zoological Garden in 2021. Her troop consists of 677 Japanese macaque monkeys who live in a sanctuary that was established in 1952 at the zoological garden. At age nine, she overthrew the dominant males in her troop and displaced her high-ranking mother as well. She became the first female leader of the troop during its recorded history of seventy years.Tumin, Remy, If you like sports, soap operas or primatology, this story is for you, Evening Briefing, The New York Times, January 21, 2022 Yakei has retained her leadership position through her first breeding season that had been thought to be a time when she might have been challenged successfully.Wartik, Nancy; Wolfe, Jonathan, The rein of Japan's monkey queen has just begun, Evening Briefing, Number 8, The New York Times, Thursday, April 7, 2022 Both scientific and popular interest is leading to extensive coverage of Yakei's behavior.


Mating and parenting
A male and female macaque form a and mate, feed, rest, and travel together during the mating season, and on average, this relationship typically lasts 16 days. Females enter into consortships with an average of four males a season. Higher-ranking males have longer consortships than their subordinates. In addition, higher-ranking males try to disrupt consortships of lower-ranking males. Females may choose to mate with males of any rank. However, dominant males mate more frequently than others, as they are more successful in mate guarding. The female decides whether mating takes place. In addition, a dominant position does not mean a male will successfully mate with a female. Males may join other troops temporarily during the mating season and mate with those females.

During the mating season, the face and genitalia of males redden and their tails stand erect,Wolfe L. (1979) "Sexual maturation among members of a transported troop of Japanese macaques". Primates 20(3):411–8. and the faces and anogenital regions of females turn scarlet. Macaques copulate both on the ground and in the trees. Roughly one in three copulations leads to .

(2009). 9781605012797, MobileReference. .
Macaques signal when they are ready to mate by looking backward over a shoulder, staying still, or walking backward toward their potential partner.Hanby JP, Brown CE. (1974) "The development of sociosexual behaviours in Japanese macaques Macaca fuscata". Behaviour 49:152–96. A female emits a "squawk", a "squeak", or produces an atonal "cackle" during copulation. Males have no copulatory vocalizations.

Females engage in same-sex mounting unrelated to the mating season and therefore, are mounted more often by other females than by males. This behavior has led to proposals in literature that female Japanese macaques are generally bisexual, rather than preferentially homo- or heterosexual.

A macaque mother moves to the periphery of her troop to give birth in a secluded spot, unless the group is moving, when the female must stay with it.Thomsen R. (1997) "Observation of periparturitional behaviour in wild Yakushima macaques ( Macaca fuscata yakui) ". Folia Primatol 68(6):338–41. Macaques usually give birth on the ground. Infants are born with dark-brown hair.Hiraiwa M (1981) "Maternal and alloparental care in a troop of free-ranging Japanese monkeys". Primates 22(3):309-29. A mother and her infant tend to avoid other troop members. The infants consume their first solid food at five to six weeks old, and by seven weeks, can forage independently from their mothers. A mother carries her infant on her belly for its first four weeks. After this time, the mother carries her infant on her back, as well. Infants continue to be carried past a year. The mother may socialize again very slowly.Bardi M, Shimizu K, Fujita S, Borgognini-Tarli S, Huffman MA. (2001) "Social behavior and hormonal correlates during the perinatal period in Japanese macaques". Horm Behav 39(3):239–46. However, has been observed, usually by females who have not had infants of their own. Male care of infants occurs in some groups, but not in others; when they do, usually, older males protect, groom, and carry an infant as a female would.Gouzoules H. (1984) "Social relations of males and infants in a troop of Japanese monkeys: a consideration of causal mechanisms". In: Taub DM, editor. Primate paternalism. New York: Van Nostrand Reinhold Co. pp. 127–45.

Infants have fully developed their locomotive abilities within three to four months.Minami T. (1974) "Early mother-infant relations in Japanese monkeys". In: Kondo S, Kawai M, Ehara A, editors. Contemporary primatology, proceedings of the 5th International Congress of Primatology. Basel(CH): S. Karger. pp. 334–340. When an infant is seven months old, its mother discourages suckling; full weaning happens by its eighteenth month.

In some populations, male infants tend to play in larger groups more often than females.Glick BB, Eaton GG, Johnson DF, Worlein J. (1986) "Social behavior of infant and mother Japanese macaques ( Macaca fuscata): effects of kinship, partner sex, and infant sex". Int J Primatol 7(2):139–55. However, female infants have more social interaction than their male counterparts, and female infants will associate with individuals of all ages and sexes. When males are two years old, they prefer to associate with other males around the same age.


Communication
During feeding or moving, Japanese macaques often emit sounds that are called "coos". These vocalizations most likely serve to keep the troop together and strengthen social relations among females.Mitani M. (1986) "Voiceprint identification and its application to sociological studies of wild Japanese monkeys" ( Macaca fuscata yakui). Primates 27(4):397-412. Macaques usually respond to coos with coos of their own.Sugiura H. (2001) "Vocal exchange of coo calls in Japanese macaques". In: Matsuzawa T, editor. Primate origins of human cognition and behavior. Tokyo: Springer. p135-54. Coos also are uttered before grooming along with vocalizations identified as "" calls. Variants of the "girney" calls are made in different contexts.Masataka N. (1989) "Motivational referents of contact calls in Japanese monkeys". Ethology 80(1-4):265-73. This call also serves as appeasement between individuals in aggressive encounters.Blount B. 1985. "Girney" vocalizations among Japanese macaque females: context and function. Primates 26(4):424-35. Macaques have alarm calls for alerting to danger and other calls to signal estrus that sound similar to danger alerts. Threat calls are heard during aggressive encounters and are often uttered by supporters of those involved in antagonistic interactions. The individual being supported supports those callers in the future.Machida S. (1990) "Threat calls in alliance formation by members of a captive group of Japanese monkeys". Primates 31(2):205-11.


Intelligence and culture
The Japanese macaque is an intelligent species. Researchers studying this species at Island in Japan left out on the beach for them to eat, then witnessed one female, named Imo (Japanese for yam or potato), washing the food off with river water rather than brushing it off as the others were doing, and later even dipping her clean food into salty seawater. Blue Planet Biomes, ¶ 12, sent. 1, downloaded 2009-02-15T16:00+09:00 Karger.com "Carrying and Washing of Grass Roots by Free-Ranging Japanese Macaques at Katsuyama" by Nakamichi, Masayuki; Kato, Eiko; Kojima, Yasuo; and Itoigawa, Naosuke in Folia Primatologica: International Journal of Primatology; Vol. 69, No. 1, 1998, § "Abstract", ¶ 1, sent. 1, downloaded 2009-02-15T16:00+09:00 After a while, other members of her troop started to copy her behavior. This trait was then passed on from generation to generation, until eventually all except the oldest members of the troop were washing their food and even seasoning it in the sea. Similarly, she was the first observed balling up wheat with air pockets and soil, throwing it all into the water, and waiting for the wheat to float back up free from the soil to consume it. An altered misaccount of this incident is the basis for the "hundredth monkey" effect.Amundson, Ron (Summer 1985). ed. "The Hundredth Monkey Phenomenon". Skeptical Inquirer: 348–356. That behavior also spread among her troop members.

The macaque has other unusual behaviours, including bathing together in hot springs and rolling snowballs for fun. In winter, the bathing is associated with lower levels of stress, with the higher ranking females dominating the restricted resource to compensate for the higher rates of stress outside of the spring. Also, in recent studies, the Japanese macaque has been found to develop different accents, similar to human cultures. National Geographic "Monkeys Have Accent, Japanese Study Finds" Macaques in areas separated by only a few hundred miles may have very different pitches in their calls, their form of communication. The Japanese macaque has been involved in many studies concerning neuroscience and also is used in drug testing.


Ecology
The Japanese macaque is . In colder areas, from autumn to early winter, macaques feed in between different activities. In the winter, macaques have two to four feeding bouts each day, with fewer daily activities. In the spring and summer, they have two or three bouts of feeding daily.Yotsumoto N. (1976) "The daily activity rhythm in a troop of wild Japanese monkey". Primates 17(2):183-204. In warmer areas such as , daily activities are more varied. The typical day for a macaque is 20.9% inactive, 22.8% traveling, 23.5% feeding, 27.9% social grooming, 1.2% self-grooming, and 3.7% other activities.Maruhashi T. (1981) "Activity patterns of a troop of Japanese monkeys ( Macaca fuscata yakui) on Yakushima island, Japan". Primates 22(1):1-14. Macaques usually sleep in trees, but they also sleep on the ground, as well as on or near rocks and fallen trees. During the winter, macaques huddle together for warmth on sleeping grounds.Takahashi H. (1997) "Huddling relationships in night-sleeping groups among wild Japanese macaques in Kinkazan island during winter". Primates 38(1):57-68. Macaques at Jigokudani Monkey Park are notable for visiting the hot springs in the winter to warm up after being encouraged to concentrate there in the 1960s, part of a plan to reduce local crop damage from foraging.


Diet
The Japanese macaque is and eats a variety of foods. More than 213 species of plants are included in the macaque's diet.
(1974). 9783805521659, S. Karger.
They also eat insects, bark, and soil, as well as flowers and some nectar. On , fruit, mature leaves, and fallen seeds are primarily eaten. The macaque also eats fungi, ferns, invertebrates, and other parts of plants. In addition, in Yakushima, their diets vary seasonally with fruits being eaten in the summer and herbs being eaten in the winter. Farther north, macaques mostly eat seasonal foods such as fruit and nuts to store fat for the winter, when food is scarce. On the northern island of , macaques mostly eat fallen seeds, herbs, young leaves, and fruits, especially of the Japanese barberry ( Berberis thunbergii) and Japanese zelkova ( ). When preferred food items are not available, macaques dig up underground plant parts (roots or rhizomes) or eat soil and fish.
(2025). 9784431538868, Springer, Tokyo.

File:Macaca fuscata yakui eating Yakiimo - 3.jpg|A macaque eating yakiimo File:Artis Japanse Makaak (36377275012).jpg|A Japanese macaque eating various fruits and vegetables


Distribution and habitat
The Japanese macaque is the northernmost-living non-human primate. It is found on three of the four main Japanese islands, south of the Blakiston's Line: , , and . The northernmost populations live on the Shimokita Peninsula, the northernmost point of Honshu.Uehara S. (1975) "The importance of the temperate forest elements among woody food plants utilized by Japanese monkeys and its possible historical meaning for the establishment of the monkeys & apes; range: a preliminary report". In: Kondo S, Kawai M, Ehara A, editors. Contemporary primatology, proceedings of the 5th International Congress of Primatology. Basel(CH):S. Karger. p392-400. Several of Japan's smaller islands are inhabited by macaques as well. The southernmost population living on is a subspecies of the mainland macaques, M. fuscata yakui. A study in 1989 estimated the total population of wild Japanese macaques to be 114,431 individuals.

The Japanese macaque lives in a variety of habitats. It inhabits subtropical forests in the southern part of its range and subarctic forests in mountainous areas in the northern part of its range. It can be found in both warm and cool forests, such as the deciduous forests of central and northern Japan and the broadleaf evergreen forests in the southwest of the islands. Warm temperate evergreen and broadleaf forests and cool temperate deciduous broadleaf forests are the most important habitats for macaques.

In 1972, a troop of approximately 150 Japanese macaques were relocated from Kyoto to a primate observatory in southwest , . The observatory is an enclosed ranch-style environment and the macaques have been allowed to roam with minimal human interference. At first, many perished in the unfamiliar habitat, which consists of arid brushland. The macaques eventually adapted to the environment, learned to avoid predators (such as eagles, , and ), and they learned to forage for mesquite beans, cactus fruits, and other foods. The surviving macaques flourished, and by 1995, the troop consisted of 500 to 600 individuals. In 1996, hunters maimed or killed four escaped macaques; as a result, legal restrictions were publicly clarified and funds were raised to establish a new sanctuary near Dilley, Texas. In 1999, the Animal Protection Institute took over management of the sanctuary and began to rescue other species of primates. As of 2017, the troop cohabitated with six other species of macaque.


Relationship with humans
Traditional human behaviors that are threats to macaques have been , use of forest woods for construction and fuel, and hunting. Since World War II, these threats have declined due to social and economic changes in Japan,Sprague DS. (2002) "Monkeys in the backyard: encroaching wildlife and rural communities in Japan". In: Fuentes A, Wolfe LD, editors. Primates face face: conservation implications of human-nonhuman primate interconnections. Cambridge (UK): Cambridge U Pr. p254-72. including the prohibition of macaque hunting in 1947.
(2025). 9781461439677, Springer Science & Business Media.
New threats have emerged, in particular the replacement of natural forests with lumber plantations.

Protection for the macaques, increased afforestation, and the human-caused extinction of their natural predators the , have led to the macaque population growing heavily since the 1940s. Because of this, and land-use changes increasing the proximity of agriculture to the macaques' range, they have become a major agricultural pest; they can climb over regular fences and quickly realise that deterrents such as scarecrows do not pose an actual threat, so methods such as electric fences must be used.渡邊邦夫 「ニホンザル」『動物世界遺産 レッド・データ・アニマルズ1 ユーラシア、北アメリカ』小原秀雄・浦本昌紀・太田英利・松井正文編著、講談社、2000年、139頁。加藤陸奥雄、沼田眞、渡辺景隆、畑正憲監修 『日本の天然記念物』、講談社、1995年、716-720頁。 In 2019, the cost of agricultural damage caused by macaques was around 900 million yen. Over 20,000 macaques are culled each year in an attempt to reduce agricultural damage, and there are concerns that this culling is reducing the macaques' range.

Macaques have often entered urban areas, with one macaque recorded living in central Tokyo for several months.Fukuda F. (2004) "Dispersal and environmental disturbance in Japanese macaques ( Macaca fuscata) ". Prim Rep 68:53-69. In 2022 the city of Yamaguchi experienced aggression from the monkeys with at least 50 people attacked.


Cultural depictions
The Japanese macaque has featured prominently in the religion, folklore, and art of Japan, as well as in proverbs and idiomatic expressions in the Japanese language.

In belief, mythical beasts known as raijū sometimes appeared as monkeys and kept , the god of lightning, company. The "three wise monkeys", who warn people to "see no evil, hear no evil and speak no evil", are carved in relief over the door of the famous Tōshō-gū shrine in Nikkō.

The Japanese macaque is a feature of several fairy tales, such as the tale of Momotarō and the fable about The Crab and the Monkey.

The monkey is part of the . That zodiac has been used for centuries in Japan and led to many representations of the macaque for that figure.

The creature was sometimes portrayed in paintings of the rich cultural epoch, the that flourished from 1603 to 1867, as a tangible metaphor for a particular year. The early nineteenth-century artist and , (1793–1841), created a painting of a macaque.

(2025). 9780231138260, Columbia University Press.
The last great master of the genre of woodblock printing and painting, Tsukioka Yoshitoshi, also featured the macaques in his prints. Also during the Edo period, numerous clasps for or pouches (collectively called ) were carved in the shape of macaques.

Spoken references to macaques abound in the history of Japan. Originating from before his rise to power, the famed samurai, Toyotomi Hideyoshi, was compared to a monkey in appearance and nicknamed Kozaru ("Little Monkey").

(1989). 069102846X, Princeton University Press. . 069102846X
In modern Japanese culture, because monkeys are considered to indulge their openly and frequently (much the same way as are thought to in some Western cultures), a man who is preoccupied with sex might be compared to or metaphorically referred to as a monkey, as might a romantically involved couple who are exceptionally amorous.


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