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Macaca sinicatoque macaque

By Matthew Kanelos

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. (Fooden, 1979)

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. (Fooden, 1979)

  • Range elevation
    0 to 2100 m
    0.00 to 6889.76 ft
  • Average elevation
    1000 m
    3280.84 ft

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. (Fooden, 1979; Harris, 1970; Napier, 1967)

  • Sexual Dimorphism
  • male larger
  • sexes colored or patterned differently
  • Range mass
    2.5 to 6.12 kg
    5.51 to 13.48 lb
  • Average mass
    4.2 kg
    9.25 lb
  • Range length
    367 to 495 mm
    14.45 to 19.49 in
  • Average length
    448 mm
    17.64 in

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. (Fooden, 1979; Michael and Crook, 1973)

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. (Fooden, 1979; Michael and Crook, 1973; Tuttle, 1975)

  • 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
  • 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

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. (Fooden, 1979; Harris, 1970; Michael and Crook, 1973)

  • 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

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. (Fooden, 1979; Michael and Crook, 1973)

  • 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

Behavior

Toque macaques are mainly arboreal, spending much of their time in trees. In trees they use a quadripedal style of locomotion. When walking on the ground, they walk on their digits. Bipedal walking occurs on the ground when the hands are full. They are social, forming groups of up to forty members. These groups are long-standing, but emigration between groups occurs in males during the late juvenile stage. In groups there is a 2 to 1 ratio of females to males. Toque macaques are very active and forage for food for a large part of every day. Males and females play different roles in groups. Males lead the group and settle fights between younger members. They play no role in the raising of young. Females are the primary care providers for young and a female's position in the dominance hierarchy may influence that of her young. The age distribution is even, with half of group members being adults and half infants or juveniles. There is a clear dominance hierarchy in groups, with the oldest male being most dominant. Following the oldest males are subadult males, adult females, and finally juveniles. (Dittus and Ratnayeke, 1989; Napier, 1967; Dittus and Ratnayeke, 1989; Napier, 1967; Dittus and Ratnayeke, 1989; Napier, 1967; Dittus and Ratnayeke, 1989; Napier, 1967; Dittus and Ratnayeke, 1989; Fooden, 1979; Napier, 1967)

  • Range territory size
    .17 to 2 km^2
  • Average territory size
    .4 km^2

Home Range

Home range sizes of toque macaques are not reported, although groups defend areas against other social groups. (Fooden, 1979)

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. (Eimerl and DeVore, 1965; Fooden, 1979)

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. (Fooden, 1979; Harris, 1970; Fooden, 1979; Harris, 1970)

  • Animal Foods
  • birds
  • mammals
  • reptiles
  • insects
  • Plant Foods
  • leaves
  • fruit
  • flowers
  • sap or other plant fluids
  • Other Foods
  • fungus

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. (Fooden, 1979)

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. (Dewit, et al., 1991; Fooden, 1979)

  • Ecosystem Impact
  • disperses seeds
Mutualist Species
Commensal/Parasitic Species

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. ("The IUCN Red List of Threatened Species", 2009)

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. ("The IUCN Red List of Threatened Species", 2009; Dewit, et al., 1991)

  • Negative Impacts
  • injures humans
    • carries human disease
  • crop pest

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. ("The IUCN Red List of Threatened Species", 2009; "U.S. Fish and Wildlife Service Threatened & Endangered Species System", 1976)

Contributors

Matthew Kanelos (author), University of Michigan-Ann Arbor, Phil Myers (editor, instructor), Museum of Zoology, University of Michigan-Ann Arbor, Tanya Dewey (editor), Animal Diversity Web.

Glossary

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

arboreal

Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

diurnal
  1. active during the day, 2. lasting for one day.
dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

fertilization

union of egg and spermatozoan

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

island endemic

animals that live only on an island or set of islands.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

omnivore

an animal that mainly eats all kinds of things, including plants and animals

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

pet trade

the business of buying and selling animals for people to keep in their homes as pets.

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

riparian

Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).

scrub forest

scrub forests develop in areas that experience dry seasons.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

swamp

a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.

tactile

uses touch to communicate

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

threatened

The term is used in the 1994 IUCN Red List of Threatened Animals to refer collectively to species categorized as Endangered (E), Vulnerable (V), Rare (R), Indeterminate (I), or Insufficiently Known (K) and in the 1996 IUCN Red List of Threatened Animals to refer collectively to species categorized as Critically Endangered (CR), Endangered (EN), or Vulnerable (VU).

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

References

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

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.

Dewit, I., W. Dittus, D. Gibson, E. Harris, J. Vercruysse. 1991. Gastro-intestinal Helminths in a Natural Population of Macaca sinica and Presbhtis spp. at Polonnaruwa, Sri Lanka. Primates, 32(3): 391-395.

Dittus, W., S. Ratnayeke. 1989. Individual and Social Behavioral responses to Injury in Wild Toque Macaques (Macaca sinica). International journal of Primatology, 10/3: 215-235.

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

Fooden, J. 1979. Taxonmy and Evolution of the sinica Group of Macaques: I. Species and Subspecies Accounts of Macaca sinica. Primates, 20(1): 109-140.

Harris, R. 1970. Feeding and Nutrition of Nonhuman Primates. New York, London: Academic Press.

Michael, R., J. Crook. 1973. Comparative Ecology and Behaviour of Primates. New York: Academic Press.

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

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

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