dcsimg

Lifespan, longevity, and ageing

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Maximum longevity: 39 years (captivity) Observations: One wild born male lived 37.1 years in captivity, making it as old as 39 when he died (Richard Weigl 2005).
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Trophic Strategy

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Long-tailed macaques are omnivores, and exploit many different food types, reflecting the diversity of habitats they can utilize. The average length of feeding bouts is 18.3 minutes. There may be on average of twenty bouts per day. They eat a wide variety of foods such as fruits, crabs, flowers, insects, leaves, fungi, grasses, and clay. Clay may be eaten for the potassium found in it, although the potassium levels in the clay are low. However, 96% of the feeding time per day is spent eating fruit. Some limited observations suggest that long-tailed macaques select fruit based on ripeness, which is based on color.

Animal Foods: insects; aquatic crustaceans

Plant Foods: leaves; fruit; flowers

Other Foods: fungus

Primary Diet: omnivore

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Associations

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The predators of this species have not been reported.

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Morphology

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The body fur of long-tailed macaques tends to be grey-brown to reddish brown. These colors are always paler ventrally. The face is brownish-grey with cheek whiskers. The eyes are directed forward for binocular vision. The nose is flat and the nostrils are narrow and close together (catarrhine condition). Long-tailed macaques have shovel-shaped incisors, conspicuous canines, and bilophodont molars. The tooth formula is I 2/2, C 1/1, PM 2/2, and M 3/3.

The body length, not including the tail, is 40 to 47 cm. The greyish- brown or reddish colored tail is 50 to 60 cm. Long-tailed macaques exhibit sexual dimorphism in size. The average weight for males is 4.8 to 7 kg and 3 to 4 kg for females, approximately 69% of average male weight.

Range mass: 3 to 7 kg.

Range length: 40 to 47 cm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Life Expectancy

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Although the lifespan of this species has not been reported, it is likely to resemble that of other members of the genus, which seem able to obtain an maximum age of around 30 years in captivity.

Average lifespan
Sex: female
Status: captivity:
25.0 years.

Average lifespan
Status: captivity:
38.0 years.

Average lifespan
Status: captivity:
37.1 years.

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Habitat

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Long-tailed macaques are "ecologically diverse." Some of the habitats in which they have been found are primary forests, disturbed and secondary forests, and riverine and coastal forests of nipa palm and mangrove. Long-tailed macaques live most successfully in disturbed habitats and on the periphery of forests.

In Sumatra, they achieve their highest population densities in mixed mangrove swamps, secondary hill forests, and riverine forests. Some were also observed in freshwater swamps, scrub grassland, lowland primary forests, and rubber groves.

In Thailand, long-tailed macaques occur in evergreen forests, bamboo forests, and in deciduous forests.

In Malaysia, they are abundant in coastal lowland forests.

This species has been observed drinking much water and eating crabs, they often live near bodies of water. Of the various habitats occupied by long-tailed macaques, the swamp forests seem to have the highest density of them.

In recent years, habitat alteration has expanded the range of some populations of long-tailed macaques. In Malaysia, cleared land, such as plantation areas, has been colonized by this species. It has been observed that some disturbed habitats have higher troop and population sizes than some pristine forests.

This species has the highest degree of arboreality of all macaque species. One study of long-tailed macaque behavior reported that they never came to the ground except within 5 m of the edge of a river near their tree. The population densities of this diurnal species vary from 10 to 400 per squared kilometer.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: forest ; rainforest ; scrub forest

Aquatic Biomes: coastal ; brackish water

Wetlands: swamp

Other Habitat Features: agricultural ; riparian

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Distribution

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Long tailed or crab eating macaques (Macaca fascicularis) are found in southeast Asia from Burma to the Philippines and southward through Indochina, Malaysia, and Indonesia. They are found as far east as the Timor Islands.

Biogeographic Regions: oriental (Native )

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Associations

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To the extent that these animals are prey for other species, they may affect populations of those species. Macaca fascicularis may be an important predator in its ecosystem, and may have some impact on prey species.

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Benefits

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Long-tailed macaques, along with other species of macaques, have benefited humans through their use as research models in immunology, surgery, toxicology, and pharmacology. They are also important members of ecosystems and may serve as a basis for ecotourism ventures. They are sometimes still hunted for food.

Positive Impacts: food ; ecotourism ; research and education

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Benefits

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Long-tailed macaques have been known to feed in cultivated fields on such items as young dry rice, cassava leaves, rubber fruit, taro plants, and other crops. They also take food from graveyards, garbage cans, and garbage pits. They have also become involved in aggresive interactions with people.

Negative Impacts: injures humans (carries human disease); crop pest

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Conservation Status

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There are nine national parks, nine reserves, and two sanctuaries in which some long-tailed macaques reside. Regardless of the type of habitat, there must be at least 500 squared kilometers of habitat necessary to support a viable population of 5,000 long-tailed macaques. This is the minimum size for a reserve for this species.

Long-tailed macaques receive some protection in temple ruins in Thailand and protection and food in temples in Bali. In Malaysia, long-tailed macaques are legally protected, and they are fed and protected in urban forests and parks. In the Philippines, there is much interest in protecting this species. In Indonesia, the species is well-protected, but some of the reserves are being considered for oil drilling and harvesting. Some people in Bali, in fact, consider these primates to be sacred. This may increase the chances of their survival in these reserves. In Thailand, long-tailed macaques may be hunted, captured, or kept in captivity only under license. The export of this species is regulated by a quota system.

Although there are some sanctuaries for long-tailed macaques, hunting is still a problem. In Thailand and Borneo, they are hunted for food. This species is also killed because it is a pest to agriculture. The fact that these macaques destroy crops has prevented some governments from making serious conservation efforts. Long-tailed macaques are collected for medical research. They are one of the five most used primate species in medical research. Many of these macaques were exported to the United States and Great Britian. Habitat loss in these organisms is occurring due to extensive logging operations.

US Federal List: no special status

CITES: appendix ii

IUCN Red List of Threatened Species: least concern

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Behavior

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As in other species of the genus Macaca, it is likely that these animals use a combination of visual (facial expression, body postures), auditory (vocalizations), physical (grooming, play, mating, aggression) and possible chemical (olfactory) signals as a part of their repertoire of communication.

Communication Channels: visual ; tactile ; acoustic ; chemical

Perception Channels: visual ; tactile ; acoustic ; chemical

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Untitled

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The long-tailed macaque is also called the crab-eating macaque, or the cynomolgus or 'java' monkey.

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Reproduction

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In Sumatra, each social group of these monkeys contains an average of 5.7 males and 9.9 adult females. Although females outnumber males, indicating that males have more mates available to them than do females, both sexes may mate with multiple partners. However, the alpha male, or highest ranking male, of the group has the greatest access to mates of any males, and probably sires the most offspring.

Mating System: polygynous ; polygynandrous (promiscuous)

Males reach sexual maturity at approximately 6 years of age, whereas females mature by about 4 years of age. Higher-ranking females become reproductively mature before lower-ranking females. The offspring of these higher-ranking females have a greater chance of survival than do the offspring of lower-ranking females. Both of these phenomena are related to the greater food availability and decreased amount of aggression experienced by the higher ranking females.

Female long-tailed macaques show a conspicuous cyclicity of sexual behavior during their menstrual cycle. As they approach ovulation, females experience a swelling of the skin in the perineal region. However, there has been no direct correlation between the frequency of copulation and the degree of swelling of genital tissues. This concealment of ovulation could exist in order to persuade consorting males to stay with females longer.

As a side effect of the prolonged estrus period and concealment of ovulation, it is difficult for humans to estimate the length of gestation. However, it seems likely that the gestation averages approximately 162 days. The average interbirth intervasl is approximately 390 days, indicating that females can produce young each year if high ranking, and every other year otherwise.

Young are nursed until they are approximately 420 days of age. The average interbirth interval is reported to be 390 days, which is less than the average time until weaning. Both the length of nursing and the interbirth interval may be affected by the rank of the mother.

It is known that the births in this species peak from May to July, corresponding with the rainy season.

Breeding interval: Females have an average interbirth interval of 390 days.

Average number of offspring: 1.

Average gestation period: 162 days.

Average weaning age: 420 days.

Average age at sexual or reproductive maturity (female): 4 years.

Average age at sexual or reproductive maturity (male): 6 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; viviparous

Average birth mass: 320 g.

Average gestation period: 165 days.

Average number of offspring: 1.

Average age at sexual or reproductive maturity (male)
Sex: male:
1544 days.

Average age at sexual or reproductive maturity (female)
Sex: female:
1238 days.

As in all primates, there is an extended period of juvenile dependence. Females provide the bulk of the care for offspring. The young receive nourishment, encouragement, grooming, protection, and teaching from the mother. Young females are likely to remain in the troup into which they were born, whereas males are likely to emmigrate at or near sexual maturity.

Parental Investment: altricial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); post-independence association with parents; extended period of juvenile learning; inherits maternal/paternal territory; maternal position in the dominance hierarchy affects status of young

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Bonadio, C. 2000. "Macaca fascicularis" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_fascicularis.html
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Christopher Bonadio, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Crab-eating macaque

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The crab-eating macaque (Macaca fascicularis), also known as the long-tailed macaque and referred to as the cynomolgus monkey in laboratories,[1] is a cercopithecine primate native to Southeast Asia. A species of macaque, the crab-eating macaque has a long history alongside humans.[6] The species has been alternately seen as an agricultural pest,[7] a sacred animal,[8] and, more recently, the subject of medical experiments.[6]

The crab-eating macaque lives in matrilineal social groups of up to eight individuals dominated by females.[9] Male members leave the group when they reach puberty.[10] It is an opportunistic omnivore[11] and has been documented using tools to obtain food in Thailand and Myanmar.[12] The crab-eating macaque is a known invasive species and a threat to biodiversity in several locations, including Hong Kong and western New Guinea.[1] The significant overlap in macaque and human living space has resulted in greater habitat loss,[6] synanthropic living, and inter- and intraspecies conflicts over resources.

Etymology

Macaca comes from the Portuguese word macaco, which was derived from makaku, a word in Ibinda, a language of Central Africa (kaku means monkey in Ibinda).[13] The specific epithet fascicularis is Latin for a small band or stripe. Sir Thomas Raffles, who gave the animal its scientific name in 1821, did not specify what he meant by the use of this word.

In Indonesia and Malaysia, the crab-eating macaque and other macaque species are known generically as kera, possibly because of their high-pitched cries.[14]

The crab-eating macaque has several common names. It is often referred to as the long-tailed macaque due to its tail, which is often longer than its body.[15] The name crab-eating macaque refers to its being often seen foraging beaches for crabs. Another common name for M. fascicularis is the cynomolgus monkey, from the name of a race of humans with long hair and handsome beards who used dogs for hunting according to Aristophanes of Byzantium, who seemingly derived the etymology of the word cynomolgus from the Greek κύων, cyon 'dog' (gen. cyno-s) and the verb ἀμέλγειν, amelgein 'to milk' (adj. amolg-os), by claiming that they milked female dogs.[16] This name is commonly used in laboratory settings.

Taxonomy

The 10 subspecies of M. fascicularis are:

Characteristics

A crab-eating macaque in West Sumatra.

The body length of the adult, which varies among subspecies, is 38–55 cm (15–22 in) with relatively short arms and legs. Males are considerably larger than females, weighing 5–9 kg (11–20 lb) compared to the 3–6 kg (6.6–13.2 lb) of females.[15] The tail is longer than the body, typically 40–65 cm (16–26 in), which is used for balance when they jump distances up to 5 m (16 ft).[15] The upper parts of the body are dark brown with light golden brown tips. The under parts are light grey with a dark grey/brown tail. Crab-eating macaques have backwards-directed crown hairs which sometimes form short crests on the midline. Their skin is black on their feet and ears, whereas the skin on the muzzle is a light grayish pink color. The eyelids often have prominent white markings and sometimes there are white spots on the ears. Males have a characteristic mustache and cheek whiskers, while females have only cheek whiskers. Crab-eating macaques have a cheek pouch which they use to store food while foraging. Females show no perineal swelling.[18]

Distribution and habitat

The crab-eating macaque's native range encompasses most of mainland Southeast Asia, from extreme southeastern Bangladesh south through the Malay Peninsula and Singapore, the Maritime Southeast Asia islands of Sumatra, Java, and Borneo, offshore islands, the islands of the Philippines, and the Nicobar Islands in the Bay of Bengal. This primate is a rare example of a terrestrial mammal that violates the Wallace line, being found out across the Lesser Sunda Islands.[18] It lives in a wide variety of habitats, including primary lowland rainforests, disturbed and secondary rainforests, shrubland, and riverine and coastal forests of nipa palm and mangrove. It also easily adjusts to human settlements and is considered sacred at some Hindu temples and on some small islands,[8] but as a pest around farms and villages. Typically, it prefers disturbed habitats and forest periphery.

Introduced range

The crab-eating macaque is an introduced alien species in several countries, including Hong Kong, Taiwan, West Papua, Papua New Guinea, New Britain, New Ireland, New Caledonia, Solomon Islands, Fiji, Tonga, Samoa, Nauru, Vanuatu, Pohnpei, Anggaur Island in Palau, and Mauritius.[19] This has led the Invasive Species Specialist Group of the International Union for Conservation of Nature to list the crab-eating macaque as one of the "100 of the World's Worst Invasive Alien Species".[20] In Mauritius, it is a threat for the endemic and endangered Roussea simplex, as it destroys its flowers.[21] It also hinders germination of some endemic trees by destroying most of their fruits when unripe[22] and competes with the endemic endangered Mauritian flying fox for native fruits.[23]

Where it is not a native species, particularly on island ecosystems whose species often evolved in isolation from large predators, it is a documented threat to many native species. The immunovaccine porcine zona pellucida (PZP), which causes infertility in females, is currently being tested in Hong Kong to investigate its use as potential population control.[18]

Behavior and ecology

Group living

Crab-eating macaque with injury to upper lip.
Juvenile crab-eating macaque in Borneo.
Juvenile crab-eating macaque in Ao Nang, Thailand.

Macaques live in social groups that contain three to 20 females, their offspring, and one or many males. The groups usually have fewer males than females. In social groups of macaques, a clear dominance hierarchy is seen among females. These ranks remain stable throughout the female's lifetime and also can be sustained through generations of matrilines. Females have their highest birth rates around 10 years of age and completely stop bearing young by age 24.[9]

The social groups of macaques are female-bonded, meaning the males will disperse at the time of puberty. Thus, group relatedness on average appears to be lower than compared to matrilines. More difference in relatedness occurs when comparing high-ranking lineages to lower ranking lineages, with higher-ranking individuals being more closely related to one another. Additionally, groups of dispersing males born into the same social groups display a range of relatedness, at times appearing to be brothers, while at other times appearing to be unrelated.[10]

In addition to the matrilineal dominance hierarchy, male dominance rankings also exist. Alpha males have a higher frequency of mating compared to their lower-ranking conspecifics. The increased success is due partially to his increased access to females and also due to female preference of an alpha male during periods of maximum fertility. Though females have a preference for alpha males, they do display promiscuous behavior. Through this behavior, females risk helping to rear a non-alpha offspring, yet benefit in two specific ways, both in regard to aggressive behavior. First, a decreased value is placed on one single copulation. Moreover, the risk of infanticide is decreased due to the uncertainty of paternity.[24]

Increasing group size leads to increased competition and energy spent trying to forage for resources, and in particular, food. Further, social tensions build and the prevalence of tension-reducing interactions like social grooming fall with larger groups. Thus, group living appears to be maintained solely due to the safety against predation.[25]

Crab-eating macaques sometimes form mixed species groups with other primate species, including the southern pig-tailed macaque, dusky langur and white-thighed surili.[26] They have been observed engaging in grooming with other primate species, including the southern pig-tailed macaque and leaf monkeys such as Raffles' banded langur and the dusty langur.[26]

Conflict

Group living in all species is dependent on the tolerance of other group members. In crab-eating macaques, successful social group living requires postconflict resolution. Usually, less dominant individuals lose to a higher-ranking individual when conflict arises. After the conflict has taken place, lower-ranking individuals tend to fear the winner of the conflict to a greater degree. In one study, this was seen in the ability to drink water together. Postconflict observations showed a staggered time between when the dominant individual begins to drink and the subordinate. Long-term studies reveal the gap in drinking time closes as the conflict moves further into the past.[27]

Grooming and support in conflict among primates is considered to be an act of reciprocal altruism. In crab-eating macaques, an experiment was performed in which individuals were given the opportunity to groom one another under three conditions: after being groomed by the other, after grooming the other, and without prior grooming. After grooming took place, the individual that received the grooming was much more likely to support their groomer than one that had not previously groomed that individual. These results support the reciprocal altruism theory of grooming in long-tailed macaques.[28]

Crab-eating macaques demonstrate two of the three forms of suggested postconflict behavior. In both captive and wild studies, the monkeys demonstrated reconciliation, or an affiliative interaction between former opponents, and redirection, or acting aggressively towards a third individual. Consolation was not seen in any study performed.[29]

Postconflict anxiety has been reported in crab-eating macaques that have acted as the aggressor. After a conflict within a group, the aggressor appears to scratch itself at a higher rate than before the conflict. Though the scratching behavior cannot definitely be termed as an anxious behavior, evidence suggests this is the case. An aggressor's scratching decreases significantly after reconciliation. This suggests reconciliation rather than a property of the conflict is the cause of the reduction in scratching behavior. Though these results seem counterintuitive, the anxiety of the aggressor appears to have a basis in the risks of ruining cooperative relationships with the opponent.[30]

Kin altruism and spite

Macaca fascicularis fascicularis at the Bukit Timah Nature Reserve – Singapore. Video Clip

In a study, a group of crab-eating macaques was given ownership of a food object. Unsurprisingly, adult females favored their own offspring by passively, yet preferentially, allowing them to feed on the objects they held. When juveniles were in possession of an object, mothers robbed them and acted aggressively at an increased rate towards their own offspring compared to other juveniles. These observations suggest close proximity influences behavior in ownership, as a mother's kin are closer to her on average. When given a nonfood object and two owners, one being a kin and one not, the rival will choose the older individual to attack regardless of kinship. Though the hypothesis remains that mother-juvenile relationships may facilitate social learning of ownership, the combined results clearly point to aggression towards the least-threatening individual.[31]

A study was conducted in which food was given to 11 females. They were then given a choice to share the food with kin or nonkin. The kin altruism hypothesis suggests the mothers would preferentially give food to their own offspring. Yet eight of the 11 females did not discriminate between kin and nonkin. The remaining three did, in fact, give more food to their kin. The results suggest it was not kin selection, but instead spite that fueled feeding kin preferentially. This is due to the observation that food was given to kin for a significantly longer period of time than needed. The benefit to the mother is decreased due to less food availability for herself and the cost remains great for nonkin due to not receiving food. If these results are correct, crab-eating macaques are unique in the animal kingdom, as they appear not only to behave according to the kin selection theory, but also act spitefully toward one another.[32]

Reproduction

Adult crab-eating macaque with a baby

After a gestation period of 162–193 days, the female gives birth to one infant. The infant's weight at birth is about 320 g (11 oz).[11] Infants are born with black fur which will begin to turn to a grey or reddish-brown shade (depending on the subspecies) after about three months of age.[15] This natal coat may indicate to others the status of the infant, and other group members treat infants with care and rush to their defense when distressed. Immigrant males sometimes kill infants not their own in order to shorten interbirth intervals. High-ranking females will sometimes kidnap the infants of lower-ranking females. These kidnappings can result in the death of the infants, as the other female is usually not lactating. A young juvenile stays mainly with its mother and relatives. As male juveniles get older, they become more peripheral to the group. Here they play together, forming crucial bonds that may help them when they leave their natal group. Males that emigrate with a partner are more successful than those that leave alone. Young females, though, stay with the group and become incorporated into the matriline into which they were born.[33]

Male crab-eating macaques groom females to increase the chance of mating. A female is more likely to engage in sexual activity with a male that has recently groomed her than with one that has not.[34]

Diet

Long-tailed macaque and her young eating a banana in Mauritius

Despite its name, the crab-eating macaque typically does not consume crabs; rather, it is an opportunistic omnivore, eating a variety of animals and plants. Although fruits and seeds make up 60 - 90% of its diet, it also eat leaves, flowers, roots, and bark.[11] It sometimes preys on vertebrates including bird chicks, nesting female birds, lizards, frogs, and fish, invertebrates, and bird eggs. In Indonesia, it has become a proficient swimmer and diver for crabs and other crustaceans in mangrove swamps. In Bukit Timah, Singapore its diet consists of 44% fruit, 27% animal matter, 15% flowers and other plant matter, and 14% food provided by humans.[35]

The crab-eating macaque exhibits particularly low tolerance for swallowing seeds. Despite its inability to digest seeds, many primates of similar size swallow large seeds, up to 25 mm (0.98 in), and simply defecate them whole. The crab-eating macaque, though, spits seeds out if they are larger than 3–4 mm (0.12–0.16 in). This decision to spit seeds is thought to be adaptive; it avoids filling the monkey's stomach with wasteful bulky seeds that cannot be used for energy.[36] It also can help the plants by distributing seeds to new areas: Crab-eating macaques eat durians such as Durio graveolens and D. zibethinus, and are a major seed disperser for the latter species.[37]

Although the crab-eating macaque is ecologically well-adapted and poses no threat to population stability of prey species in its native range, in areas where it is not native, it can pose a substantial threat to biodiversity.[38] Some believe the crab-eating macaque is responsible for the extinction of forest birds by threatening critical breeding areas [39] as well as eating the eggs and chicks of endangered forest birds.[18]

The crab-eating macaque can become a synanthrope, living off human resources. It feeds in cultivated fields on young dry rice, cassava leaves, rubber fruit, taro plants, coconuts, mangos, and other crops, often causing significant losses to local farmers. In villages, towns, and cities, it frequently takes food from garbage cans and refuse piles.[7] It can become unafraid of humans in these conditions, which can lead to macaques directly taking food from people, both passively and aggressively.[40][41]

Tool use

Stone tool usage by crab-eating macaques in Laem Son National Park in Thailand
A crab-eating macaque using a stone as tool

In Thailand and Myanmar, crab-eating macaques use stone tools to open nuts, oysters and other bivalves, and various types of sea snails, nerites, muricids and trochids along the Andaman sea coast and offshore islands.[12][42]

Another instance of tool use is washing and rubbing foods such as sweet potatoes, cassava roots, and papaya leaves before consumption. Crab-eating macaques either soak these foods in water or rub them through their hands as if to clean them. They also peel the sweet potatoes, using their incisors and canine teeth. Adolescents appear to acquire these behaviors by observational learning of older individuals.[43]

Relationship with humans

A crab-eating macaque with a soda can at Bukit Timah, Singapore
A crab-eating macaque living in a human vicinity, with a stolen asthma inhaler

Crab-eating macaques extensively overlap with humans across their range in Southeast Asia. Consequently, they live together in many locations. Some of these areas are associated with religious sites and local customs, such as the temples of Bali in Indonesia, Thailand, and Cambodia, while other areas are characterized by conflict as a result of habitat loss and competition over food and space.[6] Humans and crab-eating macaques have shared environments since prehistoric times, and both tend to frequent forest and river edge habitats. Crab-eating macaques are occasionally used as a food source for some indigenous forest-dwelling peoples. In Mauritius, they are captured and sold to the pharmaceutical industry, and in Angaur and Palau, they are sold as pets.[18] Macaques feed on sugarcane and other crops, affecting agriculture and livelihoods, and can be aggressive towards humans. Macaques may carry potentially fatal human diseases, including herpes B virus.[18] In Singapore, they have adapted into the urban environment.[44][45]

The macaques have also developed a reputation[46] for actively stealing items from humans (cameras, eyeglasses, hats, etc.), then refusing to return their stolen goods until given food in return.[47] This unique form of kleptoparasitism (known as "robbing and bartering") has primarily been observed in smaller groups of macaques living near Hindu temples and tourist-heavy areas, suggesting it is a learned behaviour within social groups, in response to realizing that humans would trade food for their possessions back.[48]

In scientific research

M. fascicularis is also used extensively in medical experiments, in particular those connected with neuroscience and disease. Due to their close physiology, they can share infections with humans. Some cases of concern have been an isolated event of Reston ebolavirus found in a captive-bred population shipped to the US from the Philippines, which was later found to be a strain of Ebola that has no known pathological consequences in humans, unlike the African strains.[6] Furthermore, they are a known carrier of monkey B virus (Herpesvirus simiae), a virus which has produced disease in some lab workers working mainly with rhesus macaques (M. mulatta).[6] Nafovanny, the largest facility for the captive breeding of nonhuman primates in the world, houses 30,000 macaques . Plasmodium knowlesi, which causes malaria in M. fascicularis, can also infect humans. A few cases have been documented in humans, but for how long humans have been getting infections of this malarial strain is unknown. It is, therefore, not possible to assess if this is a newly emerging health threat, or if just newly discovered due to improved malarial detection techniques.[6] Given the long history of humans and macaques living together in Southeast Asia, it is likely the latter.

The use of crab-eating macaques and other nonhuman primates in experimentation is controversial with critics charging that the experiments are cruel, unnecessary and lead to dubious findings.[49] One of the most well known examples of experiments on crab-eating macaques is the 1981 Silver Spring monkeys case.[50]

In 2014, 21,768 crab-eating macaques were imported in the United States to be used in experimentation.[51]

Conservation status

Female and juvenile crab-eating macaques at Djuanda Forest Park, West Java, Indonesia

The crab-eating macaque has the third-largest range of any primate species, behind only humans and rhesus macaques. The IUCN Red List categorizes the species as endangered,[1] and CITES lists them as Appendix II.[52] The species' IUCN status was changed in the summer of 2022 from the Least Concern classification in 2020 as a result of declining population resulting from hunting and troublesome interactions with humans, despite its wide range and ability to adapt to different habitats.[1] These interactions include the skyrocketing demand for crab-eating macaques by the medical industry during the Covid-19 pandemic,[1] and the rapid development of the landscape in Southeast Asia.[1] A 2008 review of their populations suggested a need for better monitoring of populations due to increased wild trade and rising levels of human-macaque conflict, which continue to decrease overall population levels despite the species' wide distribution.[53]

Each subspecies faces differing levels of threats, and too little information is available on some subspecies to assess their conditions. The M. f. umbrosa subspecies is likely of important biological significance and has been recommended as a candidate for protection in the Nicobar Islands, where its small, native population has been seriously fragmented,[54] and is listed as vulnerable on the IUCN Red List.[1] The Philippine long-tailed macaque (M. f. philippensis) is listed as near threatened, and M. f. condorensis is vulnerable.[1] All other subspecies are listed as data deficient and need further study;[1] although recent work is showing M. f. aurea and M. f. karimondjawae need increased protection.[54] One concern for conservation is, in areas where M. fascicularis is not native, their populations need to be monitored and managed to reduce their impact on native flora and fauna.[54]

Genome

The genome of the crab-eating macaque has been sequenced.

Clones

On 24 January 2018, scientists in China reported in the journal Cell the creation of two crab-eating macaque clones, named Zhong Zhong and Hua Hua, using the complex DNA transfer method that produced Dolly the sheep.[55][56][57][58][59] This makes Zhong Zhong and Hua Hua the first primates to be cloned using the somatic cell nuclear transfer method.

See also

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Crab-eating macaque: Brief Summary

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The crab-eating macaque (Macaca fascicularis), also known as the long-tailed macaque and referred to as the cynomolgus monkey in laboratories, is a cercopithecine primate native to Southeast Asia. A species of macaque, the crab-eating macaque has a long history alongside humans. The species has been alternately seen as an agricultural pest, a sacred animal, and, more recently, the subject of medical experiments.

The crab-eating macaque lives in matrilineal social groups of up to eight individuals dominated by females. Male members leave the group when they reach puberty. It is an opportunistic omnivore and has been documented using tools to obtain food in Thailand and Myanmar. The crab-eating macaque is a known invasive species and a threat to biodiversity in several locations, including Hong Kong and western New Guinea. The significant overlap in macaque and human living space has resulted in greater habitat loss, synanthropic living, and inter- and intraspecies conflicts over resources.

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