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.
- Other Geographic Terms
- island endemic
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.
- Habitat Regions
- tropical
- terrestrial
- Terrestrial Biomes
- forest
- scrub forest
- Wetlands
- swamp
- Other Habitat Features
- agricultural
- riparian
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.
- Other Physical Features
- endothermic
- homoiothermic
- bilateral symmetry
- Sexual Dimorphism
- male larger
- sexes colored or patterned differently
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.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- 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
-
provisioning
-
pre-weaning/fledging
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-independence
-
provisioning
- female
-
protecting
- female
-
provisioning
- 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.
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.
- Key Behaviors
- arboreal
- scansorial
- diurnal
- motile
- sedentary
- territorial
- social
- dominance hierarchies
Home Range
Home range sizes of toque macaques are not reported, although groups defend areas
against other social groups.
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.
- Other Communication Modes
- pheromones
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
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.
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
- Hanuman langurs ( Semnopithecus entellus )
- purple-faced langurs ( Trachypithecus vetulus )
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
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
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.
Additional Links
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.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- island endemic
-
animals that live only on an island or set of islands.
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- scrub forest
-
scrub forests develop in areas that experience dry seasons.
- swamp
-
a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.
- agricultural
-
living in landscapes dominated by human agriculture.
- riparian
-
Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).
- 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.
- 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.
- polygynandrous
-
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
- 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).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- viviparous
-
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
- 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.
- diurnal
-
- active during the day, 2. lasting for one day.
- motile
-
having the capacity to move from one place to another.
- sedentary
-
remains in the same area
- 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
- social
-
associates with others of its species; forms social groups.
- dominance hierarchies
-
ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- pheromones
-
chemicals released into air or water that are detected by and responded to by other animals of the same species
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- pet trade
-
the business of buying and selling animals for people to keep in their homes as pets.
- 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).
- herbivore
-
An animal that eats mainly plants or parts of plants.
- frugivore
-
an animal that mainly eats fruit
- omnivore
-
an animal that mainly eats all kinds of things, including plants and animals
References
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.
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 .