Overview

Brief Summary

Biology

The toque macaque is active during the day and is equally at home in the trees and on the ground (2). It feeds on a wide range of fruit, leaves, crops and insects (2), and food is stuffed into the cheek pouches with both hands to be chewed later (6). Macaques live in large groups of eight – 43 (average 20 – 25) with several adult males and females (3). Females tend to remain in the group they were born into, but males tend to disperse into new groups when they reach maturity (2) (5). Within groups, there is a strict dominance hierarchy; lower ranking individuals are ousted from good foraging areas and are less likely to survive to reproductive age than high-ranking individuals (5). Breeding tends to take place in autumn, and females give birth to a single offspring after a gestation period of around five months (5) (2). The newborn young has a soft black natal coat in the first two months of life (2).
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Comprehensive Description

Description

The toque macaque is a stocky monkey, characterised by an unusual whorl of hair on top of the head that grows from the central crown (2). The name toque macaque refers to this 'cap' of hair; toques are brimless hats. The coat is reddish or yellowish brown in colour and the underparts are paler (2). All individuals have cheek pouches, used to carry food whilst foraging (5), short, stout limbs and long tails (6). The skin on the face is naked; the ears, border of the eyelids and lower lip are pigmented black, with the remainder of the face pinkish in males, reddish in females (3). Males tend to be larger in size than females (2).
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Distribution

Range Description

The species as a whole is endemic to Sri Lanka. The subspecies M. s. sinica is found in the north and east of the island, M. s. aurifrons in the south-west, and M. s. opisthomelas in the central part. The species and the subspecies are very fragmented in their distribution, with the wet zone form, M. s. opisthomelas being highly restricted to less than 500 km2 in extent of occurrence and less than 100 km2 in area of occupancy (Molur et al. 2003).
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Geographic Range

Macaca sinica is endemic only to the island of Sri Lanka. It is found in all areas of this country excluding the Jaffina Peninsula in northern Sri Lanka and is also not found in the Trincomalee area in northeastern Sri Lanka.

Biogeographic Regions: oriental (Native )

Other Geographic Terms: island endemic

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Historic Range:
Sri Lanka

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Range

This macaque is restricted to Sri Lanka (1).
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Physical Description

Morphology

Physical Description

Toque macaques are the smallest species of Macaca. They have a golden brown coat on their dorsal surface and white hair on their ventral surface. This white coat extends to the cheeks and around the ears. The have a small tuft of hair on the top of their head that resembles a hat. The amount of hair on the top of the head is geographically variable within their range. They have a long tail that is black dorsally and pale white ventrally. Toque macaques have muscular cheek pouches that are used for storage during foraging. These pockets are lined with mucous and the muscles serve to push the food back into their mouth. Individuals have thirty two teeth; their dental formula is 2/2, 1/1, 2/2, 3/3. There is sexual dimorphism in canine size, with males having larger canines. All of their molars are quadrucuspid. Their faces are hairless. Toque macaque males have a tan face while females have different shades of pink. Males are larger than females and complete their development later. Females have a head and body length of 400 mm while males are around 475 mm. Both males and females have long tails that are slightly longer than their head and body length. Their ears and lips are black. Toque macaques may be the smallest species of Macaca, but they have one of the largest tails compared to body size. They also have very large ears that are 9% of the head and body length. Their basal metabolic requirement is 150 kcal/kg daily. There are two described subspecies of M. sinica: Macaca sinica aurifrons, which lives in the lowland dry region in northern Sri Lanka, and Macaca sinica sinica, which lives in the wet evergreen forests in southern Sri Lanka. Macaca sinica sinica is slightly larger has darker, denser, dorsal hair.

Range mass: 2.5 to 6.12 kg.

Average mass: 4.2 kg.

Range length: 367 to 495 mm.

Average length: 448 mm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger; sexes colored or patterned differently

  • Napier, J. 1967. A Handbook of Living Primates. New York: Academic Press.
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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
This species is found in a variety of forest types (Molur et al. 2003) at all altitudes up to about 2,100 m (Corbet and Hill 1992). M. s. aurifrons is found in lowland and midland tropical rainforest and wet zone lowland forests, M. s. opisthomelas in montane tropical rainforest, and M. s. sinica in dry evergreen forest near water (Molur et al. 2003). It is mainly arboreal and diurnal (Molur et al. 2003). It is frugivorous, but also consumes flowers and insects, and has been known to raid crops and garbage dumps (Molur et al. 2003).

Systems
  • Terrestrial
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Habitat

Toque macaques live in a variety of biomes throughout Sri Lanka. They spend a large amount of time in trees and live in all types of forests. The type of trees and tree height varies with region. The most important resource is a source of drinking water. This accounts for the lack of any M. sinica in the northern penninsula and parts of southern Sri Lanka. As a result of occupying separate regions two sub species are recognized. The lowland species, Macaca sinica sinica, lives in northern Sri Lanka. This region has much smaller trees and only receives one meter of rainfall a year. The highland species, Macaca sinica aurifrons, lives in the evergreen forests of southwestern Sri Lanka. This region gets over three meters of rainfall a year.

Range elevation: 0 to 2100 m.

Average elevation: 1000 m.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: forest ; scrub forest

Wetlands: swamp

Other Habitat Features: agricultural ; riparian

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Habitat

Occurs in gallery and semi-deciduous forests, scrub and by the edges of water courses, from lowlands to 1,500 metres elevation (2) (3).
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Trophic Strategy

Food Habits

Toque macaques spend much of their day foraging for food in trees and on the ground. They have special pouches in their cheeks that are used to store food during foraging. If they sense danger while foraging they will put their food in their mouth and flee to safety. Once safe they will eat the contents. The diet consists mostly of fruit. They also eat tree flowers, buds, and leaves. In their range there are 46 different types of trees, only 6 of which are not utilized for food. When available they will eat small invertebrates or vertebrates such as birds or lizards. Although these are not a main source of energy for M. sinica they represent an important source of protein and a considerable portion of time is spent searching for such prey. During the dry season water obtained from food is not enough to sustain toque macaques, so they must make daily trips to watering holes. Toque macaques also raid crop, such as rice, cocoa, and coconut.

Animal Foods: birds; mammals; reptiles; insects

Plant Foods: leaves; fruit; flowers; sap or other plant fluids

Other Foods: fungus

Primary Diet: herbivore (Frugivore ); omnivore

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Associations

Ecosystem Roles

Toque macaques play many roles in their ecosystem. It is estimated that a troop will eat up to 1 percent of the annual fruit production of the forest they live in. Through their frugivory they also help to disperse seeds. They are predators of small lizards and birds, but these are not a staple of their diet. Toque macaques a commensal relationship with two species of monkeys, Hanuman langurs (Semnopithecus entellus) and purple-faced langurs (Trachypithecus vetulus), with which they co-occur. They are often observed foraging together. They do not compete for food resources as M. sinica primarily eats fruit and both Presbytis species eat leaves. They are hosts to a variety of parasites.

Ecosystem Impact: disperses seeds

Mutualist Species:

Commensal/Parasitic Species:

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Predation

Toque macaques spend most of their time in trees, which limits their exposure to potential predators. When on the ground they appear much more cautious then when they are in the trees. They avoid open spaces when possible. When they must be in the open, they stay in compact groups. When any member senses danger the whole group will flee to nearby trees. When frightened they sometimes freeze in place when in dense foliage. To avoid predation they sleep high in forks of trees that are far from the central trunk. Groups of M. sinica rarely sleep in the same tree over consecutive nights to avoid predation. They do fall prey to large, arboreal predators, including snakes, and possibly to large avian predators, although none are reported.

Known Predators:

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Life History and Behavior

Behavior

Communication and Perception

Toque macaques are diurnal primates that rely heavily on vision. They have excellent stereoscopic and color vision. Often they distinguish between food sources using their vision, instead of smell. As in other primates, Toque macaques have excellent control over their hands and feet. They have well developed thumb-index finger control. They use acoustic communication among individuals and groups. There has been 30 different vocalizations recorded. They use warning calls to alarm other group members of danger and other vocal communication during play. There is a clear dominance hierarchy in groups of M. sinica and the dominant male is easy to identify. He is generally well groomed and muscular. Grooming is a common social activity. During the breeding season females communicate that they are in estrous by secreting a pungent mucous from their cervix that males smell.

Communication Channels: visual ; tactile ; acoustic ; chemical

Other Communication Modes: pheromones

Perception Channels: visual ; acoustic

  • Eimerl, S., I. DeVore. 1965. The Primates. New York: Time-Life Books.
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Life Expectancy

Lifespan/Longevity

The lifespan of toque macaques in the wild is about the same as in captivity, up to 35 years. The expected lifespan in the wild is low due to high infant mortality rates. There is also significant mortality among adolescent males when they venture off to join a different troop. Once toque macaques have reached sexual maturity they will likely live to an old age.

Range lifespan

Status: wild:
30 (high) years.

Range lifespan

Status: captivity:
35 (high) years.

Typical lifespan

Status: wild:
4.5 to 4.8 years.

Average lifespan

Status: captivity:
30 years.

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Lifespan, longevity, and ageing

Maximum longevity: 29.3 years (captivity) Observations: In the wild, this species has an average life span of less than 5 years (Ronald Nowak 1999). Little is known about its longevity in captivity, but one animal lived 29.3 years (Richard Weigl 2005).
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Reproduction

Reproduction

During the late summer season every sexually mature female goes into estrus once a month for three months. Females in estrus secrete a pungent mucus from their vagina which serves as a signal of her fertility to potential mates. They are promiscuous and paternity of offspring is generally unknown. Once a pair are ready to mate they withdraw from their troop but are often followed by younger males who attempt to mate with the female after the first male. Although there is a well developed dominance hierarchy among troops of M. sinica, copulation frequency is not higher among the dominant males.

Mating System: polygynandrous (promiscuous)

Toque macaques have a three month breeding season in late summer. The exact month of breeding varies with location. In the breeding season females come into estrous once a month. Mounting is always initiated by a male. There is a six month gestation period. There are 0.69 young born per mature female yearly.

Breeding interval: Toque macaques breed once yearly.

Breeding season: Breeding season is generally between July and September.

Average number of offspring: 0.69.

Range gestation period: 5 to 6 months.

Average birth mass: 390 g.

Range weaning age: 140 to 190 days.

Average weaning age: 170 days.

Average time to independence: 2 years.

Average age at sexual or reproductive maturity (female): 5 years.

Average age at sexual or reproductive maturity (male): 7 years.

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

The majority of the parental investment is from females. Females invest significantly in gestation and lactation, during late pregnancy their energy requirements almost double. Females supervise groups of playing young together. Juveniles play in groups with other members that are the same age. Mothers provide protection to their young, but a mother will only protect its youngest child in the presence of danger. Young learn by watching older members of troops. Males provide little care to the young as there is no way to be sure of paternity.

Parental Investment: altricial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female); extended period of juvenile learning

  • Tuttle, R. 1975. Primate Functional Morphology and Evolution. Paris: Mouten Publishers.
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
EN
Endangered

Red List Criteria
A2cd

Version
3.1

Year Assessed
2008

Assessor/s
Dittus, W., Watson, A. & Molur, S.

Reviewer/s
Mittermeier, R.A. & Rylands, A.B. (Primate Red List Authority)

Justification
Although a widely distributed species, the population is inferred to have declined by more than 50% in the last 40 years (approximately 3 generations) due to habitat loss at an equal or slightly higher rate. The species is also threatened from persecution, and from exploitation for the pet industry.

History
  • 2000
    Vulnerable
  • 1996
    Lower Risk/near threatened
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Conservation Status

Toque macaques are listed as endangered according to the IUCN Red List. Their population has been decreasing steadily and total population size has been cut in half over the last 40 years. This loss is due to habitat destruction and persecution of M. sinica by humans. Although they are protected internationally, they are not protected by Sri Lankan law. There are many parks and reserves in Sri Lanka where toque macaques occur.

US Federal List: threatened

CITES: no special status

State of Michigan List: no special status

IUCN Red List of Threatened Species: endangered

  • 1976. "U.S. Fish and Wildlife Service Threatened & Endangered Species System" (On-line). U.S. Fish and Wildlife Service. Accessed April 09, 2009 at http://ecos.fws.gov/speciesProfile/SpeciesReport.do?spcode=A06N.
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Current Listing Status Summary

Status: Threatened
Date Listed: 10/19/1976
Lead Region: Foreign (Region 10) 
Where Listed:


Population detail:

Population location: entire
Listing status: T

For most current information and documents related to the conservation status and management of Macaca sinica , see its USFWS Species Profile

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Status

The toque macaque (Macaca sinica) is classified as Vulnerable (VU) on the IUCN Red List (1), and is listed under Appendix II of CITES (4). The lowland wet zone toque macaque (subspecies Macaca sinica aurifrons) and dry zone toque macaque (subspecies Macaca sinica sinica) are both classified as Endangered (EN) on the IUCN Red List (1).
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Population

Population
Studies on M. s. aurifrons suggest that the animals live in smaller troops as forests become increasingly disturbed in the wet zone, and the home range becomes much wider.

Very little is known about the population of M. s. opisthomelas, though some studies exist for M. s. sinica (Molur et al. 2003). M. s. sinica has been affected in the past by stochastic events.

Population Trend
Decreasing
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Threats

Threats

Major Threats
The chief threat to this species is habitat loss owing to the encroachment of plantations, and fuel wood collection. Other threats include shooting, snaring and poisoning of the animals, as they are considered to be crop pests (Molur et al. 2003).

According to government data, during one 42-year period (1956-1993), the country lost 50% of its forest cover and more than 50% if the last 10 years (1994-2003) are included. There is a 1:1 relationship between loss of critical habitat and population number. Therefore, the species is reduced numerically minimally by 50% (Molur et al. 2003). Much of the original forested habitat of M. s. aurifrons in the southwest rainforest areas has been converted to agriculture, home gardens and plantations. These habitats are inimical to macaque survival because the animals are not tolerated by humans (Molur et al. 2003). In addition, 80% of hill country forests were lost to tea plantations during the 19th century. M. s. opisthomelas has been reduced numerically by >80% over 200 years. This trend is continuing as high elevation natural forests are being converted to agriculture (vegetable plots and dairy pasture) (Molur et al. 2003) The Mahaweli Development Scheme has destroyed much dry-zone forest habitat of M. s. sinica (Molur et al. 2003).

M. s. aurifrons and M. s. sinica are kept as pets, which is a threat in the dry zone (Nekaris pers. comm.). The animals are increasingly being used as target practice by the Sri Lankan and Tamil armies (Santiapillai and Wijeymohan 2003).
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Threats

The main threat affecting this species is habitat loss and damage as a result of wood extraction (1). In some areas they are caught for the local pet trade (6).
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Management

Conservation Actions

Conservation Actions
Although protected internationally under CITES Appendix II, this is the only endemic species not protected by law in Sri Lanka (Molur et al. 2003).

M. s. sinica and M. s. aurifrons are known to occur in numerous protected areas, including: Attidiya-Belanwila Sanctuary, Buddaragala Sanctuary, Dombagaskande Forest Reserve, Elehara Forest Reserve, Flood Plains National Park, Gal Oya National Park, Kanthale Naval Sanctuary, Kaudulla National Park, Kitulgala Sanctuary, Kurulukelle Sanctuary, Madura Oya National Park, Menikdena Archaelogical Reserve, Minneriya-Giritale National Park, Moragaswewa National Park, Muthurajawela S. Polonnaruwa Sanctuary, Remmalakanda Forest Reserve, Rendenigala Sanctuary, Ritigala Nature Reserve, Ruhuna National Park, Sinharaja Forest Reserve, Sirigiriya Sanctuary, Somawathie National Park, Thangamalai Sanctuary, Udawalawe National Park, Udawattekele Sanctuary, Wasgamuwa National Park, Wilpattu National Park, Yala National Park (Sri Lanka). M. s. opisthomelas is not found in any protected areas, though it formerly occurred in Horton Plains National Park.
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Conservation

The toque macaque is not protected by Sri Lankan law. Conservation education programmes are required to encourage the protection of the species (6).
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

There is significant persecution of toque macaques as they are considered crop pests. They are shot and poisoned as ways to keep them out of crops. They are also known zoonotic vectors of Trichuris, Ascaris, and certain strongyle worms.

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

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Economic Importance for Humans: Positive

Toque macaques are not economically important in Sri Lanka. They are widely viewed as pests because they raid crops and garbage dumps. They are sometimes sold as pets.

Positive Impacts: pet trade

  • 2009. "The IUCN Red List of Threatened Species" (On-line). Accessed April 06, 2009 at http://www.iucnredlist.org/search.
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Wikipedia

Toque macaque

The toque macaque (Macaca sinica) is a reddish-brown coloured Old World monkey endemic to Sri Lanka, where it is locally known as the 'Rilewa' or 'Rilawa' (hence 'Rillow' in the Oxford English Dictionary). It is named for the toque shaped whorl of hair on its head, rather like the bonnet of the related Bonnet Macaque.

It lives in troops, sometimes numbering up to 20, and has developed into three subspecies. It has a head and body length of 35–55 cm (14–22 in), a tail length of 40–60 cm (16–24 in) and can weigh up to 8.4 kg (19 lb).

Troops of the toque macaque are a common sight in The Cultural Triangle, where many ancient temples are situated, hence earning them the nickname of "Temple Monkey".

There are two subspecies of Toque Macaque that have been described:

References

  1. ^ Groves, C. (2005). Wilson, D. E.; Reeder, D. M. eds. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. pp. 164. OCLC 62265494. ISBN 0-801-88221-4. http://www.bucknell.edu/msw3/browse.asp?id=12100562. 
  2. ^ Dittus, W., Watson, A. & Molur, S. (2008). Macaca sinica. In: IUCN 2008. IUCN Red List of Threatened Species. Downloaded on 4 January 2009.
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