Long-tailed macaque

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TAXONOMY

Suborder: Haplorrhini
Infraorder: Simiiformes
Superfamily: Cercopithecoidea
Family: Cercopithecidae
Subfamily: Cercopithecinae
Genus: Macaca
Species: M. fascicularis
Subspecies: M. f. atriceps, M. f. aurea, M. f. condorensis, M. f. fascicularis, M. f. fusca, M. f. karimondjawae, M. f. lasiae, M. f. philippinensis, M. f. tua, M. f. umbrosa

Other names: M. cynomolgus or M. irus; crab-eating macaque, cynomolgus monkey, kera macaque, or longtail macaque; macaque crabier or macaque de buffon (French); macaca cangrejera (Spanish); javaapa or krabbmakak (Swedish); M. f. atriceps: dark-crowned long-tailed macaque; M. f. aurea: Burmese long-tailed macaque; M. f. condorensis: Con Song long-tailed macaque; M. f. fusca : Simeulue long-tailed macaque; M. f. karimondjawae: Kemujan long-tailed macaque; M. f. lasiae: Lasia long-tailed macaque; M. f. philippinensis: Philippine long-tailed macaque; M. f. tua: Maratua long-tailed macaque; M. f. umbrosa: Nicobar long-tailed macaque

Conservation status: please search the IUCN Red List.

Life span: 31 years
Total population: Unknown
Regions: Philippines, Malaysia, Indonesia, Burma, India, Vietnam, Cambodia, Laos, Thailand
Gestation: 5.5 months (165 days)
Height: 412 to 648 mm (M), 385 to 503 mm (F)
Weight: 4.7 to 8.3 kg (M), 2.5 to 5.7 kg (F)

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MORPHOLOGY

The subspecies of long-tailed macaques vary in color from light brown or grayish to brown fur covering their backs, legs, and arms while their undersides are much lighter (Rowe 1996; Groves 2001). They have pinkish-brown faces and the fur on their heads sweeps back over their forehead, often creating a crest of hair on the top of their heads. Males have mustaches and cheek whiskers that frame their faces while females have beards as well as cheek whiskers. Both males and females have white coloration on the eyelids near the nose (Rowe 1996). The defining characteristic, for which they are named, is their extraordinarily long tail that is almost always longer than their height from head to rump, and ranges in length between 400 and 655 mm (1.31 and 2.15 ft) (Fa 1989; Groves 2001). Infants have a natal coat and are born black, changing to the adult pelage as they mature (Rowe 1996). By two to three months of age, they begin to lose their black coat and by one year, they have the adult coloration (Fooden 1995). Like other macaques (Macaca), long-tailed macaques are sexually dimorphic (Dittus 2004). Males measure between 412 and 648 mm (1.35 and 2.13 ft) and weigh, on average, between 4.7 and 8.3 kg (10.4 and 18.3 lb). Females measure only 385 to 503 mm (1.26 to 1.74 ft) and have average weights between 2.5 and 5.7 kg (5.5 and 12.6 lb) (Fa 1989). In addition to being taller and heavier, males have much larger canine teeth than females (Dittus 2004). Macaques have cheek pouches in which they can store food as they forage, and transport it away from the foraging site to eat (Lucas & Corlett 1998).

Long-tailed macaque on forest floor — *Macaca fascicularis*

These macaques are primarily arboreal and can leap distances between trees up to five meters (16.4 ft), using their long tails for balance (Rodman 1991; Rowe 1996). Long-tailed macaques move quadrupedally through the canopy and spend some amount of time on the ground (Rodman 1991).

RANGE

CURRENT RANGE MAPS (IUCN REDLIST):
Macaca fascicularis

Long-tailed macaques are widespread throughout the islands of Southeast Asia into mainland Asia. They are found in the Philippines, in mainland Malaysia and on Borneo, in Indonesia on Sumatra, Java, Timor, and the Lesser Sunda Islands, in Burma, in India on the Nicobar Islands, and in Vietnam, Cambodia, Laos and Thailand (Fittinghoff & Lindburg 1980; Groves 2001). Longtail macaques are also found on Angaur Island, Palau where they were introduced in the 1900s. During the period of German rule, miners are said to have brought a pair of long-tailed macaques to the island the entire population is thought to have arisen from these two animals (Poirier & Smith 1974). They were also introduced to Tinjil Island, Indonesia in 1988 as part of the establishment of a natural habitat breeding facility to supply biomedical research (Fittinghoff & Lindburg 1980; Kyes 1993). These breeding colonies were established after a 1981 meeting of the World Health Organization which resulted in the promotion of natural breeding colonies in habitat countries that would serve as supply populations for essential biomedical research while conserving other populations of the same species (Kyes et al. 1998). Prior to the introductions in Palau and Indonesia, long-tailed macaques were introduced by Portuguese explorers to the island of Mauritius in the Indian Ocean. This introduction is thought to have occurred at least as early as the 17th century, but could have been as early as the 16th century. This population has since grown to about 35,000 animals and these long-tailed macaques are culled and exported for use in biomedical research (Sussman & Tattersall 1986; Bonnotte 2001).

HABITAT

Long-tailed macaques live in primary, secondary, coastal, mangrove, swamp, and riverine forests from sea level up to elevations of 2000 m (6561 ft) (Rowe 1996; Supriatna et al. 1996). They prefer forested areas near water and are found in higher densities near riverbanks, lakeshores, or along the seacoast (van Schaik et al. 1996). They preferentially utilize secondary forest, especially if it borders human settlement, where they have access to gardens and farms to crop-raid (Crockett & Wilson 1980; Sussman & Tattersall 1986).

Long-tailed macaques are found in tropical rain forests characterized by warm, humid climate and heavy seasonal rainfall (Supriatna et al. 1996; Umapathy et al. 2003). The rainy season in Southeast Asia lasts from about September to May, with average monthly rainfall between 140 and 300 mm (5.5 and 11.8 in) and with less rainfall from June through August or September (Lucas & Corlett 1991; Yeager 1996; Umapathy et al. 2003). Annual rainfall ranges between 2100 and 4500 mm (6.89 to 14.8 ft) per year (Cannon & Leighton 1994; Melisch & Dirgayusa 1996).

ECOLOGY

Long-tailed macaque capturing frog — *Macaca fascicularis*

Long-tailed macaques are frugivorous, and on Borneo in Kalimantan, 66.7% of their diet consists of ripe, fleshy fruits while macaques on Sumatra devote an even higher percentage of their diets to fruit (82%) (Yeager 1996; Wich et al. 2002). During times of year when fruit is unavailable, the dry season into the early rainy season, long-tailed macaques focus on other food sources including insects, stems, young and mature leaves, flowers, seeds, grass, mushrooms, invertebrates, bird eggs, clay and bark (Wheatley 1980; Yeager 1996; Son 2003). Where they forage in mangroves, long-tailed macaques spend time consuming crabs and have also been seen eating frogs, shrimp and octopus (Sussman & Tattersall 1986; Son 2003). Most of the time spent looking for and eating food is done below 20 m (65.6 ft) and usually closer to 12 m (39.4 ft) in the lower canopy, understory, and on the ground (Ungar 1996). One reason long-tailed macaques forage at this height is to avoid feeding competition with other sympatric primates. When they inhabit disturbed areas near human settlement, long-tailed macaques quickly learn to raid gardens or crops and beg for food from humans (Lucas & Corlett 1991). They have also been known to enter houses and steal food if humans are not there to frighten or deter them (Gurmaya et al. 1994). One management strategy to decrease the chance of long-tailed macaques becoming agricultural pests is for park officials to feed them and to allow tourists to feed them (Son 2004). In some parts of their range where long-tailed macaques come into contact with tourists at nature reserves up to 22% of their diet can be from provisioned foods (Lucas & Corlett 1991; Son 2003). This strategy is not without costs to both macaques and humans. When humans feed long-tailed macaques, both contact and non-contact aggression increases within and between groups of macaques at the same feeding site. Serious injury or death can occur because of aggressive encounters between long-tailed macaques (Wheatley 1991). Humans are also at risk when feeding long-tailed macaques because the possibility of disease transmission is increased when the two species come into contact. Humans who feed and interact with the macaques can be scratched or bitten and exposed to zoonotic diseases such as simian foamy virus, a virus in the same family as HIV (Jones-Engel et al. 2005). Furthermore, in addition to viruses that have been identified, there is the potential for previously unknown diseases to transfer from the monkeys to the humans who are feeding them, posing a serious health risk to the surrounding communities and beyond (Jones-Engel et al. 2005).

The daily time budget and movement patterns of long-tailed macaques involve traveling, feeding, resting and socializing. Long-tailed macaques have a home range size of about 1.25 km² (.58 mi²) and daily path length varies greatly between 150 and 1900 m (.09 and 1.18 mi) (Wheatley 1980; Rowe 1996). They highly prefer foraging and moving in riverine habitats, and the amount of time spent foraging decreases as they move further from the riverbank. Most of their daily activity happens within 100 m (328 ft) of the river, where resource density is much higher than areas further inland (van Schaik et al. 1996). After leaving their nesting tree between 5:30 and 6:00 a.m., they spend the morning hours feeding heavily while they spend the midday hours resting (Gurmaya et al. 1994). Aggressive interactions between group members are the highest while feeding on fruit, indicating strong competition for this valuable food resource (van Schaik & van Noordwijk 1988). While the group is resting, individuals sleep, play, or groom, with the youngest animals spending time playing while the older animals sleep and groom (Son 2004). After the midday period of rest, long-tailed macaques continue to search for food and feed as they move closer to their sleeping trees. They enter the sleeping trees in the early evening, between 6:00 and 6:30 p.m. and stay there for the night (Gurmaya et al. 1994; Son 2004).

Long-tailed macaque swimming — *Macaca fascicularis*

Long-tailed macaques sleep in trees along the river and are particular about choosing their roosting sites. Each group sleeps in its own tree and individuals huddle together when they sleep to maintain body temperature. They sleep toward the edge of the branches near the top or crown of the tree and preferentially choose branches that overhang the river (van Schaik et al. 1996). Long-tailed macaques are excellent swimmers, and this may be a predator avoidance technique: if they are threatened, they simply can escape by dropping to the water and swimming to safety (Rowe 1996; van Schaik et al. 1996). Some predators include pythons, monitor lizards, raptors, large cats, and, in some areas, feral dogs. (Palombit 1992; van Noordwijk & van Schaik 1999). The felid predation risk is so strong in some parts of their range that there is a discernible effect on social structure and group size (van Schaik & van Noordwijk 1985).

Content last modified: January 6, 2006

Written by Kristina Cawthon Lang. Reviewed by Carolyn Crockett.

Cite this page as:
Cawthon Lang KA. 2006 January 6. Primate Factsheets: Long-tailed macaque (Macaca fascicularis) Taxonomy, Morphology, & Ecology . <http://pin.primate.wisc.edu/factsheets/entry/long-tailed_macaque/taxon>. Accessed 2020 July 17.

SOCIAL ORGANIZATION AND BEHAVIOR

Long-tailed macaque group grooming — *Macaca fascicularis*

Long-tailed macaques live in multi-male/multi-female groups of six to 58 individuals, with the smallest groups found where there are no felids on the island of Simeulue in Indonesia (van Schaik & van Noordwijk 1985). One reason primates form groups is the benefit of increased protection against predators. In a group, there is higher likelihood of detecting a predator, and the chance of an individual primate becoming a victim of that predator decreases as group size increases. Frugivorous primates living in areas with predators must make a tradeoff; increasing group size enhances protection against predators but it also increases competition for fruit resources, which can be densely clumped and seasonally variable (van Schaik & van Noordwijk 1988; Sterck & Steenbeek 1997). On Simeulue, long-tailed macaques do not have to maintain large groups because of the lack of predators and therefore may have smaller groups to avoid feeding competition (van Schaik & van Noordwijk 1985). One way to reduce feeding competition in larger groups during times of fruit scarcity in areas other than Simeulue is for long-tailed macaque groups to split up into smaller foraging parties (Wheatley 1980; van Noordwijk & van Schaik 1987; van Schaik & van Noordwijk 1988).

Female long-tailed macaques remain in their natal groups and exhibit strong dominance hierarchies in which rank is passed on from mother to daughter and remains within a matriline (de Jong et al. 1994; van Noordwijk & van Schaik 1999). Females in a long-tailed macaque group are related in some way, either as sisters, half-sisters, cousins, or mother-daughter (de Jong et al. 1994). One measure of rank, especially among females, is the direction of grooming. Higher-ranking individuals enjoy more and longer-lasting grooming sessions from low-ranking individuals than vice versa (Wheatley 1999). Other important indicators of rank among female long-tailed macaques are the “bared-teeth display,” in which the submissive female bares her teeth to the dominant female, and displacement, in which a dominant female displaces a submissive animal at a feeding site (van Noordwijk & van Schaik 1987; Sterck & Steenbeek 1997). When long-tailed macaque groups divide and forage separately, the main group consists of the highest-ranking females while the lower-ranking females form their own subgroup to forage (Sterck & Steenbeek 1997). High-ranking females benefit because of easier access to food, increased safety from predators and aggressive male macaques, as well as increased reproductive success (Wheatley 1999).

Males also exhibit a strict dominance hierarchy, with the highest-ranking male having the highest access to reproductive females and fathering the majority of infants born in the group during his tenure (Engelhardt et al. 2004). The second-ranking male, or beta male, fathers the remaining 20% of infants born into the group (de Jong et al. 1994). Aggressive interactions between males result in serious injuries, especially lacerations from their long, sharp canine teeth. Injuries obtained while fighting can lead to increased mortality through infection or predation (Wheatley 1999; van Noordwijk & van Schaik 2001). The relationship between the alpha female and male is maintained by grooming. The alpha female grooms the alpha male frequently, and interferes through aggressive behavior if lower-ranking females associate with him. Males of any rank groom females most frequently during estrus (Wheatley 1999).

Long-tailed macaque group inspecting injured male — *Macaca fascicularis*

Males emigrate from their natal groups with groups of their peers before sexual maturity, usually between four and six years old (de Jong et al. 1994). They generally immigrate into adjacent groups, but in their new groups young males will not attempt to establish themselves in the dominance hierarchy until at least seven years of age. Usually by nine years, they will attempt to take over the top position. If one male of the peer group attains the top rank, rather than challenging him, his peers will immigrate into a new group (van Noordwijk & van Schaik 2001). Males migrate multiple times over their lives and if they are unsuccessful at achieving the top dominance rank, they will immigrate into a new group and attempt another take-over. The average residence of males in a group is 45 months (van Noordwijk & van Schaik 1999; 2001). Even though they leave their natal groups, the sons of high-ranking females are more likely to have higher ranks in the dominance hierarchy of their new group than sons of low-ranking females (van Noordwijk & van Schaik 1999). Once a young male gains the position of highest-ranking male in the group, he will only hold that position, on average, for a period of three years (de Jong et al. 1994). When a male takes over for a former dominant male, the group conditions are conducive to infanticide. Males kill infants that are unlikely to be their own in order to shorten interbirth intervals. Females that lose a nursing infant will come into estrus faster than if they reared the infant completely. By inducing cycling in females, infanticidal males thus increase their chances of siring offspring as soon as possible after a take-over (Hrdy 1974).

REPRODUCTION

Reproductive output is linked to dominance status among female long-tailed macaques. The highest-ranking females have more offspring over their lifetimes than lower-ranking females, not only because they begin to reproduce at a younger age but also because their offspring have a higher chance of survival (van Noordwijk & van Schaik 1999). Some factors that contribute to the higher rate of survival for high-ranking females’ offspring include: better access to food, a more central position in the main foraging group which offers protected from predators and a decreased likelihood of being harassed by low-ranking females (van Noordwijk & van Schaik 1987). Long-tailed macaques exhibit birth seasonality, which varies slightly across their range. In Gunung Leuser National Park on Sumatra, the majority of the births occur between July and November (van Noordwijk & van Schaik 1999). Where they are studied on West Java, there is a concentration of births in January and February (Engelhardt et al. 2004). Female reproductive success is linked to food availability; during years of high food abundance, birth rates are higher than years with food scarcity and in years with mast fruiting events, births occur earlier and are more frequent than in years with average fruit availability (van Noordwijk & van Schaik 1987; van Schaik & van Noordwijk 1988).

Long-tailed macaque mother and infant — *Macaca fascicularis*

Sexual maturity in females is reached at four years of age and high-ranking daughters begin reproducing before 5.5 years of age while low-ranking daughters begin reproducing after 5.5 years. Males reach sexual maturity by age seven (Varavudhi et al. 1992; de Jong et al. 1994). Estrus is characterized by sex skin swelling, the inflation of the skin surrounding the anogenital region, and behavior patterns (Engelhardt et al. 2005). Females have a distinct set of vocalizations referred to as “copulation calls” that are heard during 80% of the copulations. Female long-tailed macaques mate multiple times throughout the day during the period of fertility (Engelhardt et al. 2004). About 80% of the infants born into a long-tailed macaque group are sired by the alpha male. One of the ways he ensures he is the dominant breeder is to guard the receptive females from subordinate males in the group (de Jong et al 1994; Engelhardt et al. 2004). The dominant male remains within five meters (16.4 ft) of the sexually receptive female and prevents other males from mating with her by aggressively displaying and physically preventing them from inspecting or coming near her (Engelhardt et al. 2004). Another male will take an opportunity to attempt to mate with the female if the dominant male is not closely guarding her, but the alpha male usually replaces him through aggressive behavior such as chasing, biting, or lunging (Engelhardt et al. 2005). Mate guarding is an energetically expensive behavior and is only seen in the few days surrounding a female’s peak period of receptivity. Male long-tailed macaques are able to discern this period of receptivity because of behaviors exhibited by the female, which include solicitation, copulation calls, and grooming after copulation as well as apparent pheremonal cues (Engelhardt et al. 2004).

Females give birth to singletons and the interbirth interval averages 18 months, with females more likely to skip a year after giving birth to a surviving infant than after a year without successful reproduction (van Schaik & van Noordwijk 1988; de Jon et al. 1994). Females reach peak reproductive capacity at age 10 and they continue to reproduce until about 24 years of age, though reproduction significantly decreases after age 20 (van Noordwijk & van Schaik 1999).

PARENTAL CARE

Long-tailed macaque mothers are the primary caregivers of their newborn infants and are very protective of them, not allowing infants out of their grasp (Wheatley 1999). During the first weeks of life, the mother long-tailed macaque maintains contact with her infant almost constantly, but as the infant ages the amount of contact decreases (Meishvili et al. 1991). By four months, the mother spends significantly less time in contact with the infant and begins to exhibit normal ovarian cycles again. By the time the infant is 10 or 11 months of age the mother sharp rejects it and may act aggressively if it tries to nurse. (Meishvili et al. 1991).

Long-tailed macaque adult and infant — *Macaca fascicularis*

Other females in the group are intensely interested in newborns, and attempt to touch, groom, and take the infant from the mother. Sometimes they are successful in “kidnapping” the infant, and if the mother is unable to retrieve the newborn, it is likely to die. High-ranking females that “kidnap” low-ranking infants are more likely to prevent the mother from rescuing the infant and subsequently the infant is more likely to die (Wheatley 1999). If an infant dies, the mother will not reproduce again until the next year, thus “kidnapping” may be a way for high-ranking females to decrease the reproductive success of low-ranking females in the group. In the first year of life, the survival rate of long-tailed macaque infants is 81%, but the overall survival rate of long-tailed macaques from birth to four years is 68% (van Noordwijk & van Schaik 1999). When infants reach juvenilehood, adult males spend more time playing with them, soliciting juveniles by making certain vocalizations (Wheatley 1999).

COMMUNICATION

Long-tailed macaques have an extensive vocal communication repertoire for use in many situations. There are two general classes of long-tailed macaque vocalizations, “harsh” and “clear” calls. Some important “harsh calls” are the “kra call,” named for its sound, that are performed by allage and sex classes and are used in both mildly and highly excited macaques to show alarm and are named for their sound; “alarm calls,” given by long-tailed macaques in the presence of a potential predator or when they feel threatened and are made of three to five, chirp-like pulses and “barks” which are heard during aggressive interactions between individuals (Palombit 1992; Wheatley 1999). “Clear calls” include a variety of “coos,” which promote friendly interactions and avoid aggression between individuals and are heard between subordinate and dominant females as well as infants calling to their mothers (Wheatley 1999). Other calls include “screams” which are calls heard in aggressive interactions, especially by subordinates that are being victimized, “affiliation calls” given by females in an attempt to get closer to another female’s infant, and “geckers” heard from infants during weaning conflicts or when in other situations of conflict (Wheatley 1999).

Content last modified: January 6, 2006

Written by Kristina Cawthon Lang. Reviewed by Carolyn Crockett.

Cite this page as:
Cawthon Lang KA. 2006 January 6. Primate Factsheets: Long-tailed macaque (Macaca fascicularis) Behavior . <http://pin.primate.wisc.edu/factsheets/entry/long-tailed_macaque/behav>. Accessed 2020 July 17.

INTERNATIONAL STATUS

For individual primate species conservation status, please search the IUCN Red List.
Also search the current scientific literature for primate conservation status (overall as well as for individual species), and visit CITES (Convention on International Trade in Endangered Species of Wild Fauna and Flora).

Conservation information last updated in 2006 follows, for comparison:

CONSERVATION THREATS & POTENTIAL SOLUTIONS

Threat: Invasive Alien Species

Though there are no data available for the effects of invasive alien species on long-tailed macaque conservation, the long-tailed macaque is itself an invasive alien species causing serious damage to parts of its range where it has been introduced and populations grow unchecked due to a lack of natural predators (Poirier & Smith 1974; Crockett pers. comm.). For example, the population of long-tailed macaques established on the island of Mauritius has created serious conservation concerns for other endemic species. With no natural predators on the island, the long-tailed macaque population has grown unchecked for almost a half century (Stanley 2003). They consume seeds of native plants and help spread exotic weeds throughout the forests, changing the composition of native forests, causing them to be quickly overrun by invasive plant species. They also have directly contributed to the near extinction of several bird species on Mauritius, including the Mauritian green parrot and the pink pigeon. Long-tailed macaques destroy the nests of these birds as they move through their home ranges and cause direct mortality by eating the eggs of these critically endangered species (Stanley 2003; Temple pers. comm.).

Threat: Persecution

Though they are not currently threatened, long-tailed macaques are subject to persecution where they come into contact with humans as crop raiders. They can be serious agricultural pests, raiding gardens and eating fruits including oranges, bananas, and coconuts as well as staple crops such as taro, sweet potatoes, rice, and cassava root. This relationship has probably been going on since human agriculture began in the range of long-tailed macaques. In response, humans often shoot the monkeys and can eliminate entire local populations (Wheatley et al. 1999).

Threat: Human Disturbance

Harvesting long-tailed macaques for biomedical research was once a serious threat to survival but this practice has largely ended (Supriatna et al. 1996). A natural habitat breeding colony in Indonesia was established in the 1980s in order to harvest from semi-wild, managed populations instead of natural populations (Kyes et al. 1998).

LINKS TO MORE ABOUT CONSERVATION

CONSERVATION INFORMATION

No current links for Macaca fascicularis
Links for all species

CONSERVATION NEWS

Conservationists sound alarm over macaque (AFP; July 15, 2011)
Impact of the Japan earthquake and tsunami on animals and environment (Scientific American; March 22, 2011)
Monkey business puts macaques in danger (Sydney Morning Herald; October 3, 2007)
Monkey export ban lifted (The Star, Malaysia; August 18, 2007)
Malaysia seizes 900 monkeys from poachers (Reuters; July 9, 2007)
Links for all species

ORGANIZATIONS INVOLVED IN Macaca fascicularis CONSERVATION

Content last modified: January 6, 2006

Written by Kristina Cawthon Lang. Reviewed by Carolyn Crockett.

Cite this page as:
Cawthon Lang KA. 2006 January 6. Primate Factsheets: Long-tailed macaque (Macaca fascicularis) Conservation . <http://pin.primate.wisc.edu/factsheets/entry/long-tailed_macaque/cons>. Accessed 2020 July 17.

The following references were used in the writing of this factsheet. To find current references for Macaca fascicularis, search PrimateLit.

REFERENCES

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Crockett CM, Wilson WL. 1980. The ecological separation of Macaca nemestrina and M. fascicularis in Sumatra. In: Lindburg DG, editor. The macaques: studies in ecology, behavior and evolution. New York: Van Nostrand Reinhold. p 148-81.

de Jong G, de Ruiter JR, Haring R. 1994. Genetic structure of a population with social structure and migration. In: Loeschcke V, Tomiuk J, Jain SK, editors. WXS 68, Conservation genetics. Basel (Switzerland): Birkhäuser Verlag. p 147-64.

de Ruiter J, van Hooff JARAM, Scheffrahn W. 1994. Social and genetic aspects of paternity in wild long-tailed macaques (Macaca fascicularis). Behaviour 129(3-4): 203-24.

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Engelhardt A, Hodges JK, Niemitz C, Heistermann M. 2005. Female sexual behavior, but not sex skin swelling, reliably indicates the timing of the fertile phase in wild long-tailed macaques (Macaca fascicularis). Horm Behav 47(5): 195-204.

Engelhardt A, Pfeifer JB, Heistermann M, Niemitz C, van Hoof JARAM, Hodges JK. 2004. Assessment of female reproductive status by male longtailed macaques, Macaca fascicularis, under natural conditions. Anim Behav 67(5): 915-24.

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Fooden J. 1995. Systematic review of southeast Asian longtail macaques, Macaca fascicularis (Raffles, [1821]). Field Zool 81: 1-206.

Groves C. 2001. Primate taxonomy. Washington DC: Smithsonian Inst Pr. 350 p.

Gurmaya KJ, Adiputra IMW, Saryatiman AB, Danardono SN, Sibuea TTH. 1994. A preliminary study on ecology and conservation of the Java primates in Ujung Kulon National Park, West Java, Indonesia. In: Thierry B, Anderson JR, Roeder JJ, Herrenschmidt N, editors. Current Primatology. Volume 1, Ecology and evolution. Strasbourg (France): Univ Louis Pasteur. p 87-92.

Hrdy SB. 1974. Male-male competition and infanticide among langurs (Presbytis entellus) of Abu, Rajasthan. Folia Primatol 22(1): 19-58.

Jones-Engel L, Engel GA, Schillaci MA, Rompis A, Putra A, Suaryana KM, Fuentes A, Beer B, Hicks S, White R, et al. 2005. Primate to human retroviral transmission in Asia. Emerg Infect Dis 11(7): 1028-35.

Kyes RC. 1993. Survey of the long-tailed macaques introduced onto Tinjil Island, Indonesia. Am J Primatol 31(1): 77-83.

Kyes RC, Sajuthi D, Iskandar E, Iskandriati D, Pamungkas J, Crockett CM. 1998. Management of a natural habitat breeding colony of longtailed macaques. Tropic Biodiv 5(2): 127-37.

Lucas PW, Corlett RT. 1991. Relationship between the diet of Macaca fascicularis and forest phenology. Folia 57(4): 201-15.

Lucas PW, Corlett RT. 1998. Seed dispersal by long-tailed macaques. Am J Primatol 45(1): 29-44.

Meishvili NV, Chalyan VG, Vancatova MA. 1991. Mother-infant relationships in cynomolgus monkeys (Macaca fascicularis). Prim Rep 29: 21-6.

Melisch R, Dirgayusa IWA. 1996. Notes on the grizzled leaf monkey (Presbytis comata) from two nature reserves in West Java, Indonesia. Asian Prim 6(1-2): 5-11.

Palombit RA. 1992. A preliminary study of vocal communication in wild long-tailed macaques (Macaca fascicularis). I. Vocal repertoire and call emission. Int J Primatol 13(2): 143-82.

Poirier FE, Smith EO. 1974. The crab-eating macaques (Macaca fascicularis) of Angaur Island, Palau, Micronesia. Folia Primatol 22: 258-306.

Rodman PS. 1991. Structural differentiation of microhabitats of sympatric Macaca fascicularis and M. nemestrina in East Kalimantan, Indonesia. Int J Primatol 12(4): 357-75.

Rowe N. 1996. The pictorial guide to the living primates. East Hampton (NY): Pogonias Pr. 263 p.

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Content last modified: January 6, 2006

IMAGES

	Macaca fascicularis Photo: Anne Savage
	Macaca fascicularis Photo: Anne Zeller
	Macaca fascicularis Photo: Diana Mossman
	Macaca fascicularis Photo: John Handford
	Macaca fascicularis Photo: Roy Fontaine
	Macaca fascicularis Photo: Roy Fontaine
	Macaca fascicularis Photo: Roy Fontaine
	Macaca fascicularis Photo: Roy Fontaine
	Macaca fascicularis Photo: Roy Fontaine
	Macaca fascicularis Photo: Roy Fontaine
	Macaca fascicularis Photo: Roy Fontaine

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