Japanese macaque, Macaca fuscata
JAPANESE MACAQUE
Macaca fuscata
Geographic Distribution and Habitat
Japanese macaques, more descriptively known as Japanese snow macaques and simply referred to as snow monkeys, inhabit three of Japan’s four main islands: Honshu, Shikoku, and Kyushu (they do not occur on Hokkaido, the fourth and northernmost island). These iconic primates also inhabit the smaller islands of Awaji, Shodo, Yakushima, Kinkazan, and Kojim, as well as other smaller islands within Japan’s different prefectures. A population who once resided on the island of Tanegashima (or “Tane Island”) within Kagoshima Prefecture is now extinct there, according to the International Union for the Conservation of Nature (IUCN).
Their expansive geographic range encompasses diverse climates and habitats, a testament to the species’ adaptability. The northern limit—the northernmost tip of Honshu Island on the Shimokita peninsula—is blanketed by more than 3 feet (1 meter) of snow for several months of the year, accompanied by frigid temperatures as low as -4 degrees Fahrenheit (-20 degrees Celsius). The Japanese macaques who live here fully embody their nickname of “snow monkey.” A series of mountain ranges known as the Japanese Alps bisect Honshu Island. Steep cliffs overlook thermal springs (in which these snow monkeys soak themselves) from which steam rises, enveloping the valley. Coniferous evergreens and broad-leaved deciduous trees are the dominant vegetation. Those Japanese macaques who live in this snowy, subarctic, mountainous region represent the world’s northernmost wild populations of all non-human primates.
The southernmost limit is Yakushima Island, home to the subspecies the Yakushima macaque (M. fuscata yakui), and provides an extreme contrast with its warm temperatures and subtropical lowland forests. In the regions between these two disparate environments, both warm and cool temperate forests provide habitat. Indicative of the habitat in which they occur, these primates reside from ground level up to 4,921 feet (1,500 meters) above sea level.
Wildlife biologists agree on two subspecies:
The Hondo macaque, the nominate subspecies (Macaca fuscata fuscata), is found within the geographic range and shared habitat environments of the “parent species,” the Japanese macaque (Macaca fuscata).
The Yakushima macaque (Macaca fuscata yakui), also known as the Yaku macaque, is found only on Yakushima Island, the southernmost limit for Japanese macaques. The IUCN states that it also once inhabited Tane Island, but ambiguity clouds this statement as to whether it refers to the parent species, the Yaku macaque subspecies—or both.
Size, Weight, and Lifespan
Sexual dimorphism (distinct differences in size or appearance between the sexes, in addition to differences in their reproductive organs) is prominent in this species. Japanese macaques are considered medium-sized monkeys. Males weigh 25 pounds (11.3 kilograms) on average with a head-to-body length of 22.4 inches (57 centimeters). Females weigh 19 pounds (8.6 kilograms) on average with a head-to-body length of 20.4 inches (52 centimeters). Those Japanese macaques residing in the species’ northernmost range tend to weigh more than their southern neighbors. Their excess body weight helps these monkeys endure the frigid temperatures of their subarctic habitat.
Mother Nature has left the species with a short stump of a tail, known as a “vestigial tail,” no longer than 3.9 inches (10 centimeters) tacked onto the monkeys’ bottoms. Over the years, as the Japanese macaque evolved, its tail became shorter and shorter—not quite disappearing (as seen—or rather, not seen, with the Barbary macaque (Macaca sylvanus)) but no longer a functional appendage. Not so for its cousin, the crab-eating macaque (Macaca fascicularis), whose alternate name of long-tailed macaque is fittingly accurate with a tail length of 19-23.5 inches (40-60 centimeters) that helps with balance, though the tail is nonprehensile.
Lifespan for Japanese macaques is reported as 28 years for males and 32 years for females. Colluding negative environmental factors and anthropogenic activities, however, can result in a much shorter longevity.
Appearance
A pinkish-red to ruby-colored face, the result of visible blood vessels (capillaries) just beneath the skin, gives Japanese macaques their arresting, human-like countenance. Soulful eyes peer out from beneath a distinct brow ridge. Eye color varies among individuals, and, on occasion, is asymmetric with each eye being a different color. Sex appears to have no bearing on eye color, but significant difference occurs depending on age. Most infants have blue eyes, but by the time they reach juvenile stage, eye color usually (but not always) changes to varying shades of orange, copper, yellow, or hazel. Eye color is thought to be inherited from parents.
A thick, lush fur coat that varies in color from brown to yellow-brown, to gray frames the pink face, giving dramatic contrast, concealing the ears in adults, and covering the macaque’s entire stocky body—except for the rump, which also happens to be pink or ruby-colored, like the face. As the temperature plummets to a low of -4 degrees Fahrenheit (-20 degrees Celsius), a Japanese macaque’s fur grows thicker, an adaptation to the cold. Most primates would succumb to this extreme cold in just a few minutes, and most cannot survive in temperatures less than 23 degrees Fahrenheit (5 degrees Celsius).
A thicker fur coat in response to environmental conditions is not this monkey’s only natural transformation. The pinkness on its face and rump, for both males and females, becomes more vibrant for some individuals—but wildlife biologists are unsure about the factors that influence this bodily response. Initially, these scientific folk believed that, for females, a more intense color signaled ovulation and fertility. But close examination of coloration data for the face and rump, along with analysis of female hormone levels during mating season—debunked the fertility response theory.
The revised thinking is that a female’s vibrant red skin color acts as an indicator (or “badge”) of her social status. Furthermore, a vibrantly red rump suggests a female is of a higher social status than those females with paler pink rumps—a distinction previously reported as being exclusive to males. Apart from social ranking, wildlife biologists posit that a more vibrant color may be in response to the macaque’s emotional and physical states. Is it possible that, like us humans, a redder face might be a response to anger or agitation? (Probably not a response to embarrassment, as we can guess that nonhuman primates don’t embarrass as easily—or maybe they simply don’t engage in our embarrassing behavior.)
Hands that are nearly identical to human hands but with much shorter opposable thumbs, huge canine teeth, and large cheek pouches that act as food storage bins (while the monkeys forage) further characterize the physical appearance of Japanese macaques.
Diet
Japanese macaques are omnivores, meaning they eat both plant and animal matter. Menu items vary according to the season. Plant-based foods from over 213 species make up the bulk of their diet. Buds, leaves, fruits, flowers, seeds, and nuts are dietary staples. When these foods are scarce, the monkeys forage for fungi and use their sharp fangs to gouge and eat tree bark. But Japanese macaques also have a palate for insects, spiders, bird eggs, snails, crabs, crayfish, and fish. Their ability to adapt their diet to available food resources across diverse habitats is key to the species’ survival.
As example, individuals living in the Japanese Alps of Honshu eat aquatic insects and fish. Those individuals residing in the species’ southernmost range near human settlements raid and eat crops—making these opportunistic monkeys quite unpopular with the humans who regard the macaques as agricultural pests in need of dispatch.
Behavior and Lifestyle
These incredibly furry monkeys are diurnal, active during daylight hours. And they are mostly terrestrial, traveling and foraging on the ground. But they venture into the trees when they feel like it. Apart from choosing an overnight sleeping site, for some groups, their arboreal exploits appear to be for fun. Japanese macaques are superb jumpers, and their leaps have been likened to that of the obstacle-leaping sport of parkour. Though they don’t typically hang by their arms from tree limbs, as some monkeys do, Japanese macaques could win a medal for swimming; they are capable of swimming more than 0.3 miles (0.5 kilometers).
Females reportedly spend more time in trees than males, while males spend more time on the ground than females. The species is primarily quadrupedal (using all four limbs to travel), but Japanese macaques sometimes walk bipedally (upright on their hind legs)—particularly when carrying something, like a fish, in both hands. Their opposable thumbs not only allow them to securely grab onto branches as they advance from one tree to the next but also assist in grabbing and holding objects.
Their decision to choose trees as overnight sleeping sites or to sleep on the ground is largely determined by the presence or absence of predators. Natural predators include mountain hawk eagles (Nisaetus nipalensis), raccoon dogs (Nyctereutes viverrinus), and feral dogs. The macaques had also once been preyed upon by the Japanese wolf (Canis lupus hodophilax), but this species has been extinct for more than 100 years—or has it? (Some enthusiasts believe they have evidence that these apex predators, so-called “ghost wolves,” still roam the Japanese mountains.)
Those Japanese macaques who share habitat with predators are more likely to sleep in trees, either individually or, when the temperature drops, huddled together for warmth. They choose deciduous trees to prevent accumulated snow from falling on top of them.
When no predators are present, as is the case on the small, mountainous island of Kinkazan (also known as Kinkasan) at the tip of the Oshika Peninsula in Miyagi Prefecture, these macaques typically choose to sleep on the ground. Seasonal and environmental conditions are additional determining factors. During the spring and summer, the macaques sleep at lower altitudes in grasslands where food is abundant. Autumn finds the macaques in sleeping sites at a range of altitudes, from sea level to 984 feet (300 meters), but they avoid high-altitude forests dominated by Japanese beech trees where strong winds are prevalent. During winter months, the macaques show a preference for valleys with fallen trees and rocky ridges that offer a greater degree of shelter from the wind and cold, allowing the monkeys to conserve energy and maintain a core body temperature independent of environmental temperature. (This natural phenomenon is known as thermoregulation).
Of course, Japanese macaques are renowned for another strategy that they use to keep warm in their frigid, snowy habitat: Those residing in Honshu’s Japanese Alps, commonly referred to by their nickname of snow monkey, routinely soak themselves in geothermal springs heated by nearby volcanoes. Poofy, fluffy heads and torsos bob and splash in the steamy water amidst a wintry landscape. Bath time is a family affair, and young offspring stay close to their mothers.
But these “hot tub” visits are learned behavior. Back in the early 1960s, in the course of conducting a field study on macaque behavior, wildlife researchers tossed a handful of soybeans into a hot spring. A Japanese macaque whom the researchers named “Mukubili” waded into the hot water to retrieve this free handout. And she stayed there. The 109-degree Fahrenheit (43 degrees Celsius) water temperature warmed her body, and her children and other young monkeys joined her. Eventually, over a period of many months, all troop members were soaking in the hot spring to keep warm. Fast forward to today: these snow monkeys are iconic for their hot spring soaks.
Anthropologists had once believed that only humans passed on learned behaviors to other individuals and across generations, a process called “cultural transmission.” Snow monkeys demolished this hypothesis.
The practice of soaking in hot springs is not the only example of cultural transmission with Japanese macaques. Their practice of washing foods prior to eating is another. Researchers conducting a field study in the 1950s observed a female macaque, dubbed “Imo,” rinse the sand off a sweet potato she was about to eat by dipping the tuber in a freshwater stream. Other members of her troop followed her lead, and this behavior has since been passed down through generations.
Over time, the macaques began rinsing potatoes in salt water. This modification led researchers to add their own culinary, if not scientific, spin on the practice: By rinsing the potatoes in salt water, rather than fresh, the macaques intentionally season their food for an enhanced epicurean experience. Aquatic insects provide the macaques with snacks and bamboo leaves cleanse their palate.
One of the most remarkable evolved foraging behaviors is seen in those individuals living in the Japanese Alps of Honshu, within the remote mountainous valley of Kamikōchi, western Nagano Prefecture. These macaques have taught themselves to fish! Researchers believe the behavior is unique to this population, evolved to obtain important animal protein during winter months when other foods are unavailable. So how does a snow monkey catch a fish? With its hands, of course. By overturning rocks where fish are hiding, chasing a fish into shallow water, grabbing the fish with both hands and then chomping down on the fish with its mouth.
Unsurprisingly, scientists consider macaques to possess an enhanced general intelligence when compared with other mammals. These primates’ innovative survival behaviors offer proof.
Japanese macaques are highly mythologized in their native Japan. In the Shinto belief, they sometimes appear as mythical beasts known as raijū, who keep Raijin, the god of lightning, company.
Perhaps they are most well-known, worldwide, as the Three Wise Monkeys, who “see no evil, hear no, evil, and speak no evil.” Depicted in various artwork, carvings, and (probably!) memes, this pictorial proverb was first carved into a wooden panel over the door of the famous Tōshōgū Shrine in Nikko, Japan, by sculptor Hidari Jingoro, in the 17th century.
Highly social creatures, Japanese macaques live in multimale-multifemale groups (known in the primate world as “troops”) composed of 10, to 20, to 100 . . . or more individuals. A troop follows a strict chain of command led by a dominant male, known as the alpha, who ranks above all other members. His special privileges included mating with all the females in his troop. But his position also comes with great responsibility; it’s up to him to protect members from intruders and predators and otherwise keep everyone safe. In larger troops, several male subleaders help the alpha male keep everyone in line. The alpha male’s partner is the alpha female. She ranks above other females in the troop, as do her children who will eventually inherit her ranking. Among the males, social ranking is determined by age, with older individuals claiming a higher rung in the hierarchy. On occasion, an exceptionally aggressive (and maybe scheming) male might muscle his way up the social ladder. Sometimes, an alpha male has gained his rank because his mother was a high-ranking female.
Upon attaining sexual maturity, males leave their natal (birth) group. As they strike out on their own, they often travel between troops with the hope of one day instilling themselves as a troop’s alpha male leader. Females remain with their birth group.
Japanese macaques travel, eat, socialize, play-fight by throwing stones or snowballs (behavior associated with youngsters), sun themselves as they rest, and warm themselves and relax in hot springs (the northernmost population) during the course of a day.
The average daily home range is about 1.43 square miles (3.7 square kilometers) and correlates both to their habitat type and to the season.
Besides their natural predators (mountain hawk eagles, raccoon dogs, and feral dogs), sympatric species include the Japanese weasel, Japanese serow, Japanese squirrel, Japanese giant flying squirrel, Japanese dwarf flying squirrel, Japanese red-backed vole, the Japanese dormouse, the Japanese bear, Okinawa spiny rat, Amami rabbit, and wild boar.
On the mountainous island of Kinkazan, Japanese macaques share habitat with more than 500 sika deer, with whom the macaques have a commensal relationship. The deer compete with the macaques for several species of plants and are opportunists in their own right. When the macaques climb trees, they invariably and unintentionally drop leaves to the forest floor which the deer readily eat.
These ungulates are regarded as messengers of the gods in the Shinto religion. Indigenous people have created a shrine where visitors to the island can purchase deer feed to give the deer.
The communication system of Japanese macaques is complex. They use a variety of vocalizations that relate to their emotional or physical state, respond to happenings in their environment, and convey specific messages to one another: as example, an individual might warn fellow troop members of a predator threat. Interestingly, researchers have identified distinct accents among different troops. Facial expressions (no “poker faces” with these primates) and body postures are also used to get a message across.
These are tactile primates who routinely engage in self-grooming and allogrooming. Allogrooming is an important activity that helps to develop social bonds with one another. Females spend a lot of time grooming their young. Besides fostering the mother-child bond, babies learn to groom themselves. But grooming also serves a pragmatic reason. By removing tangles—and parasites—the macaque’s fur coats provide better insulation against the cold.
During mating season when competition is high, a virile male might use his huge canine teeth as weapons to challenge another male for the opportunity to mate with a female. Such an occurrence is not the norm, however.
Japanese macaques are polygamous, meaning both males and females take multiple partners during a single breeding season. Females tend to be picky about with whom they mate and are more impressed by those males with higher ranking. They might bicker or scream at one another if they’ve set their eyes on the same potential male partner. Notably, they avoid mating with a partner with whom they have mated in the past 4 to 5 years, a decision that helps to prevent interbreeding. Breeding occurs from autumn through winter, with most births occurring from April through July, and again May through September.
At the beginning of her pregnancy, a female’s genitals will imbue a deeper pink or ruby coloration. Apart from signaling social rank, when associated with pregnancy a deeper color in the genitals outlines complex sexual signaling in the species, say researchers. They support their claim by noting that these reproductive females seem to conceal their ovulation cycle. And yet . . . the faces of both the male and female turn a deep scarlet red during “mating season.” (What’s up with that?)
Females attain sexual maturity (the ability to conceive and bear young) at about 3-1/2 years of age. Males lag slightly behind, attaining sexual maturity (the ability to sire young) at 4-1/2 years of age. After a pregnancy of about 5-1/2 to 6 months, a female gives birth to a single infant who weighs a mere 1.1 pounds (0.5 kilograms); twins are rare in the species. Infants are entirely dependent on their mothers for their first two years of life, at which time they are considered weaned. In the interim, infants cling to their mother as she travels and forages and pays close attention to the survival lessons she teaches them. Their bond will last a lifetime. Because a reproductive female’s interbirth interval is two or three years, a mother might be caring for a newborn while simultaneously nursing her baby’s older sibling. This situation places physical and emotional stress on mothers. Fortunately, other troop members—most often females who have no offspring of their own—will relieve a mother’s burden by helping out as babysitters.
Thanks to their largely frugivorous lifestyle, Japanese macaques are an integral species in helping to regenerate their forest ecosystem. As they travel, they disperse the seeds of the many fruits they eat, via their feces, throughout their habitat—thereby encouraging new plant growth.
Japanese macaques are classified as Least Concern by the International Union for the Conservation of Nature (IUCN, November 2015), appearing on the IUCN Red List of Threatened Species. The species’ widespread geographic distribution, its residency in many protected areas, and no serious decline in overall population (estimated at more than 100,000 individuals in their natural habitats) form the rationale for this minimal threat level.
Both subspecies, the Hondo macaque (Macaca fuscata fuscata) and the Yakushima macaque (Macaca fuscata yakui), are also classified as Least Concern.
But we must not take a lackadaisical view toward protecting this iconic primate. Each year, farmers kill upward of 10,000 Japanese macaques to keep the monkeys from eating their crops. We can’t blame the macaques for foraging on agricultural tracts of land; this land had once been their natural, forested habitat. We (humans) cut it down.
Diminished habitat can lead to interbreeding with introduced macaque species, such as the rhesus macaque (M. mulatta) and the Formosan rock macaque (M. cyclopis)—consequently, compromising the species’ genetic integrity. Most of the introduced Formosan rock macaque population has been eradicated, however.
As anthropogenic activities continue to ravage and diminish the Japanese macaque habitat, the species will slide toward decline.
Biomedical research is another threat to the species, dating back to 1965 when a troop of 55 Japanese macaques were taken from their home and flown across the globe to the Oregon National Primate Research Center. Scientists were especially interested in conducting aging and longevity studies on the species, aimed at understanding the cellular and molecular changes underlying normative primate aging. Because Japanese macaques can contract many of the same diseases as humans, these primates remain favored laboratory test subjects. Today, captive Japanese Macaques are purposely bred at Japanese research facilities (under the auspices of the National Bioresource) to study the field of neuroscience.
Japanese macaques are listed in Appendix II of the Convention on International Trade in Endangered Species (CITES), an international agreement between governments whose goal is to ensure that international trade in specimens of wild animals and plants does not threaten their survival. The species has been officially protected since 1947 under Japanese law when the government issued the edict that the macaques could no longer be hunted as a game species. In 1970, the Japanese government declared the species “a natural treasure of the nation.” Unfortunately, the rights of farmers have taken precedence over these protections, allowing farmers to kill “crop-raiding” macaques.
Wildlife reserves provide the species with greater protection, and supplemental feeding programs (started by researchers interested in studying wild Japanese macaques in their natural habitat) help those macaques living in the northernmost range survive the winter.
Because the macaques’ fondness for soaking in hot springs during the winter months eventually started to annoy the local human primates (LHP) who liked to do the same—and didn’t feel like sharing bath water with monkeys—LHP built the macaques their very own private hot spring pools within Jigokudani Monkey Park in Nagano. The Japanese macaque-only pools are today a major tourist attraction; people come from all over the world to watch and photograph the monkeys. While tourism centered on watching these snow monkeys soak in hot springs helps raise awareness and appreciation for the species, it comes with challenges. Interactions between these human primates and nonhuman primates must be monitored to ensure that both species remain safe.
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Written by Kathleen Downey, October 2024