Overview

Brief Summary

Biology

Mandrills live in mixed groups of up to 40, which come together to form troops of more than 600 individuals. There is a strict hierarchy amongst the group: a dominant male, huge and vividly coloured, heads each group, mating with fertile females and fathering almost all of the infants (6). A troop moves over a range of up to 50 km², scent-marking the territory and defending it against rivals. Groups are extremely noisy, with individuals communicating with deep grunts and high pitched crowing as they feed; when it is time to move on the alpha male emits a two-phase grunt. Mandrills spend most of their day foraging for fruits and seeds, eggs and small animals, and when night falls they retire to the trees for safety (5). When the females are receptive their rumps swell and become a more intense red, signaling her reproductive status as 'in oestrus'. Females give birth to one offspring every 18 months or so. The infant clings to her belly, and when it is heavier rides on her back (5).
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Description

The mandrill is not only the largest monkey in the world, but it is also one of the most distinctive (3). It has an extremely striking face, with a red stripe down the nose and blue flanges framing it (5). The male has a much more colourful face than females and juveniles, with more prominent hair growth around the nostrils and huge canine teeth, measuring up to 6.5 cm long, which he exposes to threaten any rivals or predators (5). This amazing monkey has red fur patches above the eyes and a yellow mane-like beard, while the rest of its body is covered with thick olive green fur. Its underparts are grey, tinged with yellow, and its body is stocky, with a short tail and a brilliantly blue to purple coloured rump (6). Males are much larger than females, and more boldly coloured. With this spectacular appearance, the male mandrill declares its identity to other animals, as well as announcing his sex and virility to females (5).
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Mandrills according to MammalMAP

Mandrills (Mandrillus sphinx) are the largest - and probably most conspicuous - of all monkeys. Male mandrills are far more colourful than females, and they use this colour as an advert of their virility as they try to win over the ladies. These social primates live in large, noisy troops headed up by a dominant male (‘drill sergeant?) who reigns over the lower ranking individuals. They are known to occur only in West Africa in Cameroon, Gabon, Equatorial Guinea and Congo.

Mandrill population are buckling under the strain of habitat loss as their natural forests give way to crops and villages. They are also targeted as bushmeat and consequently under IUCN species classifications they are considered ‘Vulnerable’.

For some more information about Mandrills look here. For more information about Mammalmap visit the MammalMAP virtual museum or blog.

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Distribution

Range Description

This species is found in moist evergreen rainforest in central west Africa, south of the Sanaga River (Cameroon) through to mainland Equatorial Guinea, western Gabon, and south-western Congo (Brazzaville) to the Kouilou River, and down to the Congo River. Inland, the Ivindo River and Ogooue River in Gabon limit its distribution to the east. In Cameroon, it is not known to occur east of the Dja river. It does not occur in the forests of south-east Cameroon or east of the Congo River. Telfer et al's (2003) study indicate that the Ogooué River, Gabon, bisects the range of the two populations, seperating them into two distinct populations 1) Cameroon and northern Gabon and 2) southern Gabon.
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Historic Range:
Equatorial West Africa

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Geographic Range

Mandrills are found in southwestern Cameroon, western Gabon, Equatorial Guinea, and southwestern Congo.

Biogeographic Regions: ethiopian (Native )

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Range

This species is found in Africa; in south-western Cameroon, western Gabon, Equatorial Guinea, and south western Congo (2).
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Physical Description

Morphology

Physical Description

Mandrills reach a height of about 80 cm. The species is characterized by a large head, a compact body with long, powerful limbs, and a stubby tail, which is held upright. The wide rotating range of the clavicles enables climbing trees, the quadrupedal walk, and the functioning of the arms. Opposable thumbs allow these monkeys to grasp tree branches. Both sexes have paired mammary glands in the chest region.

The pelage is an olive green with paler underparts. It has a brilliantly colored blue to purple naked rump. A mandrill's face has a red stripe down the middle of the muzzle and aroung the nostrils, while the sides of the muzzle are ridged lengthwise and colored blue. This helps to distinguish this species from drills which have jet black faces. Mandrills have red fur patches above the eyes and a yellow beard. These colorings are duller in females and juveniles than in adult males.

These animals are reported to have average weights of 11.5 for females, and 25 kg for males. Males are significantly larger than females and may weigh up to 54 kg.

The head and body measurements range between 610 and 764 mm.

Range mass: 54 (high) kg.

Average mass: 11.5-25 kg.

Range length: 610 to 764 mm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger; male more colorful; ornamentation

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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
Mandrills are found in evergreen rainforest, stretching between 100 and 300 km inland from the Atlantic coast, as well as in montane forest, and thick secondary forest. Mandrills are known to stretch to forest fragments in savanna and have been known to go into plantations. They are semi-terrestrial and forage primarily at less than 5 m off the ground. They are omnivorous and diets are diverse, including fruits, buds, leaves, roots, insects, fungus, and seeds. Mandrills prefer fruits when they are available, although in their primary forest habitat the fruiting of trees and lianas is irregular, leading to periodic fruit shortages: when this occurs they are highly reliant on having abundant herbaceous growth to eat. When food is scarce (e.g. during and at the end of the dry season), they also raid crops from farms.
Mandrill home ranges may be 30-50 sq km.

Systems
  • Terrestrial
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Mandrills are found in tropical rain forest habitats, montane and thick secondary forests, and thick bush. Although they are adapted to live in the ground, mandrills seek shelter in the trees during the night.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: forest ; rainforest

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This striking primate inhabits rainforests and sub-tropical forests, from flat plateaus to mountainous terrain (6).
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Trophic Strategy

Food Habits

Mandrills have a highly varied diet including fruit, seeds, fungi, roots, insects, snails, worms, frogs, lizards, and sometimes snakes and even small vertebrates. Generally, mandrill males scrounge for food on the ground while females and their young sit in midlevel trees.

Animal Foods: amphibians; reptiles; insects; mollusks; terrestrial worms

Plant Foods: roots and tubers; seeds, grains, and nuts; fruit

Other Foods: fungus

Primary Diet: omnivore

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Associations

Ecosystem Roles

These monkeys are likely to play some role in seed dispersal. To the extent that they serve as predators or as prey, they may have some effect on local food webs.

Ecosystem Impact: disperses seeds

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Predation

Predators of this species have not been reported, but are likely to include large carnivores, such as leopards.

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Life History and Behavior

Behavior

Communication and Perception

As described above in the section for behavior, communication is varied and complex in this species. It involves a variety of components, including visual and accoustic signals, scents, and tactile information.

Communication Channels: visual ; tactile ; acoustic ; chemical

Other Communication Modes: scent marks

Perception Channels: visual ; acoustic

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Life Expectancy

Lifespan/Longevity

The maximum lifespan reported for this genus is 46 years.

Average lifespan

Sex: female

Status: captivity:
31.7 years.

Average lifespan

Status: captivity:
46.0 years.

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Lifespan, longevity, and ageing

Maximum longevity: 40 years (captivity) Observations: There have been claims of animals living up to 46.3 years (http://pin.primate.wisc.edu/), which is not impossible but has not been verified either. In captivity, the record longevity of this species belongs to one female that was still alive at age 40 (Richard Weigl 2005).
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Reproduction

Mandrills live in groups, mostly in a harem structure, where a dominant male defends a group of females to whom he has exclusive mating rights.

Mating System: polygynous

Breeding is not seasonal but rather occurs about every two years, depending on the available food supply. Mating is believed to occur between July and October, while birthing occurs between December and April. Females give birth to their first young anywhere between 4 and 8 years of age. Gestation lasts for about 6 months after which females give birth to a single young. Twin young have only been observed in capivity. Infants are born with a black natal coat and pink skin, both of which endure for the first two months of life. (Macdonald, 1987)

Breeding interval: Mandrills breed every two years.

Breeding season: Mating is believed to occur between July and October.

Average number of offspring: 1.

Average gestation period: 6 months.

Range age at sexual or reproductive maturity (female): 4 to 8 years.

Key Reproductive Features: iteroparous ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); viviparous

Average birth mass: 613 g.

Average gestation period: 173 days.

Average number of offspring: 1.

Parental investment has not been extensively characterized in this species. However, it is likely that these animals are similar to other primates in which breeding occurs in a harem polygyny situation.

The bulk of the care for infants in such species is provided by the mother. Mothers give their young protection, grooming, and nourishment (milk). However, aunts, sisters, cousins, and other offspring of the mother may provide some care for young, including carrying, playing with, and grooming the young.

In species where one male mates with females, males also provide parental care. This may be direct, in the form of carrying, playing with, and grooming young, or it may be indirect, in that the father protects all the members of his harem group from potentially dangerous rival males.

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

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Molecular Biology and Genetics

Molecular Biology

Barcode data: Mandrillus sphinx

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 3 barcode sequences available from BOLD and GenBank.  Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.  See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

ATTGGAACCCTATACCTACTATTTGGTGCATGAGCCGGAGTTATAGGCACAGCCCTA---AGCCTTCTCATTCGAGCTGAACTAGGTCAACCCGGTAGTCTACTAGGCAAT---GATCACATTTACAACGTTATCGTAACGGCCCATGCATTTGTCATAATTTTCTTCATAGTTATACCCATCATGATTGGAGGCTTTGGAAACTGACTAGTGCCCCTAATA---ATTGGCGCTCCTGACATAGCATTCCCTCGTTTAAATAACATAAGCTTCTGACTCCTTCCCCCTTCCTTCCTACTACTAATAGCATCAACTATACTAGAAGCCGGCGCTGGAACAGGTTGAACAGTATACCCTCCTTTAGCGGGAAACTTATCCCACCCAGGAGCCTCCGTAGACCTA---GTCATTTTTTCCCTTCACCTAGCAGGCATTTCTTCTATTTTAGGGGCCATTAACTTTATTACTACTATTATCAACATAAAGCCTCCCGCAATATCTCAGTACCAAACTCCCCTATTCGTCTGATCAATCTTAATCACAGCAATCCTTCTACTCCTCTCACTACCAGTCCTGGCCGCC---GGCATTACCATGCTATTAACAGATCGTAACCTCAACACTACCTTCTTCGATCCAATTGGAGGAGGAGATCCCATTCTATATCAACAC
-- end --

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Statistics of barcoding coverage: Mandrillus sphinx

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 3
Specimens with Barcodes: 3
Species With Barcodes: 1
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
VU
Vulnerable

Red List Criteria
A2cd

Version
3.1

Year Assessed
2008

Assessor/s
Oates, J.F. & Butynski, T.M.

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

Contributor/s

Justification
This species is listed as Vulnerable in view of the intensive hunting pressure on this species (combined with the habitat loss) across its range, which is likely to have resulted in a decline exceeding 30% over the past 30 years.

History
  • 2000
    Vulnerable
  • 1996
    Lower Risk/near threatened
    (Baillie and Groombridge 1996)
  • 1994
    Vulnerable
    (Groombridge 1994)
  • 1990
    Vulnerable
    (IUCN 1990)
  • 1988
    Vulnerable
    (IUCN Conservation Monitoring Centre 1988)
  • 1986
    Vulnerable
    (IUCN Conservation Monitoring Centre 1986)
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Current Listing Status Summary

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


Population detail:

Population location: Entire
Listing status: E

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

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There has been a drastic decline in the mandrill population during recent years due to habitat destruction. They are especially vulnerable to hunters because of their loud calls. Mandrills are hunted as a local food source in several areas. Currently, mandrills occupy forests at a very low density and are poorly protectd if at all. As a result, they may be threatened with complete extinction in the wild. (Gale, 146)

US Federal List: endangered

CITES: appendix i

IUCN Red List of Threatened Species: vulnerable

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Status

Classified as Vulnerable (VU A2cd) on the IUCN Red List 2004 (1) and listed on Appendix I of CITES (4).
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Population

Population
Total numbers of this species are unknown, but it has undoubtably declined in recent years. It is generally rare, and in some places has been locally exterminated. The largest remaining populations are probably to be found in Gabon.
In the mid-1970s, numbers in the Wonga-Wongue National Park were said to be "fair-sized". Mandrills live at relatively low densities.

Population Trend
Unknown
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Threats

Major Threats
This species is affected by the destruction of its evergreen forest habitat since this reduces the capacity of environments to support Mandrill populations. However, the most immediate threat is posed by hunting for their meat (which is highly prized in Gabon). Commercial bushmeat hunters pose a particular threat to populations which are located close to main roads and towns. Mandrills appear to be most seriously threatened in Congo (Brazzaville).
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Mandrill populations have suffered drastic declines due to hunting for their meat. Because they travel in large troops and are easily located by their constant grunting and screaming, entire populations are quickly decimated. Hunting has become lucrative and with the use of dogs, high-powered rifles, spotlights, deep-freezers and trucks it is an even greater threat today than ever before. As human settlements expand, the mandrills are losing their habitats to logging and clearing for agriculture (2).
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Management

Conservation Actions

Conservation Actions
Mandrills are listed under Appendix I of CITES, and as Class B under the African Convention. Several reserves are located within the Mandrill's range, the most important of which is Lope National Park in Gabon. Other areas containing Mandrills need immediate protection, both legal and practical, against logging and hunting. Surveys are urgently needed to determine where viable populations exist.
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Conservation

International trade is prohibited by the mandrill's listing on Appendix I of the Convention on International Trade in Endangered Species (CITES) (4). However, even if international trade is controlled, there are still substantial threats to the mandrill. Due to severely limited funds for conservation in West Africa, and the difficulties in monitoring these animals in forests, this species is poorly protected if at all. Extinction in the wild is sadly therefore a serious threat for this most spectacular primate (2).
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Positive

Mandrills are commonly found in zoos. Becuse of their long life spans, they are valuable, longtime residents. The are also hunted for their meat in some areas.

Positive Impacts: food ; research and education

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Wikipedia

Mandrill

This article is about the primate. For other uses, see Mandrill (disambiguation).

The mandrill (Mandrillus sphinx) is a primate of the Old World monkey (Cercopithecidae) family,[4] closely related to the baboons and even more closely to the drill. It is found in southern Cameroon, Gabon, Equatorial Guinea, and Congo. Mandrills mostly live in tropical rainforests and forest-savanna mosaics. They live in groups called hordes. Mandrills have an omnivorous diet consisting mostly of fruits and insects. Their mating season takes place from June to October.

Both the mandrill and the drill were once classified as baboons in genus Papio, but recent research has determined they should be separated into their own genus, Mandrillus.[4] Mandrills are the world's largest monkeys. Charles Darwin wrote in The Descent of Man that "no other member in the whole class of mammals is coloured in so extraordinary a manner as the adult male mandrills."[5] The mandrill is classified as vulnerable by IUCN.

Description[edit]

A mandrill displays his exotic coloring.
The male mandrill's exotic coloring

The mandrill is the most colorful primate. It has an olive green or dark grey pelage with yellow and black bands and a white belly. Its hairless face has an elongated muzzle with distinctive characteristics such as a red stripe down the middle and protruding blue ridges on the sides. It also has red nostrils and lips, a yellow beard and white tuffs. The areas around the genitals and the anus are multi-colored, being colored red, pink, blue, scarlet, and purple.[6] They also have pale pink ischial callosities.[6] The coloration of the animal is more pronounced in dominant adult males. Both sexes have chest glands which are used in olfactory communication. These, too, are more prominent in dominant adult males.[7] Males also have longer canines than females, with an average of 4.5 cm (1.8 in) and 1.0 cm, respectively.[8]

The mandrill has one of the greatest sexual dimorphisms among the primates.[9] Males typically weigh 19–37 kg (42–82 lb), with an average mass of 32.3 kg (71 lb). Females weigh roughly half as much as the male, at 10–15 kg (22–33 lb) and an average of 12.4 kg (27 lb).[10] Exceptionally large males can weigh up to 54 kg (119 lb).[11][12][13] The average male is 75 to 95 cm (30 to 37 in) long and the female is 55–66 cm (22–26 in), with the short tail adding another 5–10 cm (2.0–3.9 in).[14][15] The shoulder height while on all fours can range from 45 to 50 cm (18 to 20 in) in females and 55 to 65 cm (22 to 26 in) in males. The male Mandrill is the heaviest monkey in the world, although its total length is relatively short due to its vestigial tail and, due to its high sexual dimorphism, baboons such as the Chacma and Olive average around the same weight. Compared to the largest baboons, the mandrill is more ape-like in structure, with a muscular and compact build, shorter, thicker limbs that are longer in the front and almost no tail.[16][17][18] Mandrills can survive up to 31 years in captivity. Females reach sexual maturity at about 3.5 years.

Mandrill skull (Muséum de Toulouse)

Ecology and activities[edit]

Close-up of a male mandrill's colorful face

The mandrill is found in Nigeria, southern Cameroon, Gabon, Equatorial Guinea, and Congo. Its distribution is bounded by the Sanaga River to the north and the Ogooué and White Rivers to the east. Recent research suggests mandrill populations north and south of the Ogooué river are so genetically different as to be separate subspecies. Mandrills prefer to live in tropical rainforests and forest-savanna mosaics. They also live in gallery forests adjacent to savannas, as well as rocky forests, riparian forests, cultivated areas and flooded forests and stream beds.[19][20] Mandrills will cross grass areas within their forest habitats.[21][22]

The mandrill is an omnivore. It usually consumes plants, of which it eats over a hundred species. It prefers to eat fruits, but will also eat leaves, lianas, bark, stems, and fibers. It also consumes mushrooms and soil.[20] Carnivorously, mandrills mostly eat invertebrates, particularly ants, beetles, termites, crickets, spiders, snails, and scorpions. It will also eat eggs, and even vertebrates such as birds, tortoises, frogs, porcupines, rats, and shrews.[20] Mandrills likely will eat larger vertebrates when they have the opportunity, such as juvenile bay duikers and other small antelope. Large prey are likely killed with a bite to the nape with the mandrill's long canines.[23] In males, the canines can measure over 6 cm (2.4 in) in length.[16] One study found the mandrill's diet was composed of fruit (50.7%), seeds (26.0%), leaves (8.2%), pith (6.8%), flowers (2.7%), and animal foods (4.1%), with other foods making up the remaining (1.4%).[24]

Mandrills are preyed upon mainly by leopards, in addition to crowned eagles and African Rock Pythons.[21] They may be bitten and killed by Gaboon vipers when they accidentally rouse the venomous snake. It is thought that most predators are a threat mainly to young mandrills, with the likelihood of predation decreasing in adult females and especially adult males. In a study where a mandrill troop was exposed to stimuli relating to their natural predators, only the leopard caused the larger part of the group to flee into trees. However, the large, dominant males were observed to remain in response to the images of the natural predators, even the leopard, and pace back and forth whilst baring their teeth, generally indicating aggression and the defensive role they may play in such circumstances.[25]

Mandrills are mostly terrestrial but they are more arboreal than baboons and feed as high as the canopy.[6][8] When on the ground, mandrills walk by digitigrade quadrupedalism (walking on the toes of all four limbs). When in the trees, they often move by lateral jumps.[19] Mandrills are mostly diurnal, with activities extending from morning to evening.[26] They sleep in trees at a different site each night.[20] Mandrills have been observed using tools; In the wild and in captivity, mandrills have been observed using sticks to clean themselves.[27]

Social behavior and reproduction[edit]

Mandrills seem to live in large, stable groups called "hordes". Hordes often number in the hundreds, possibly averaging around 620 individuals and reaching as many as 845.[19][21][22] It is difficult to accurately estimate group size in the forest, but filming a group crossing a gap between two forest patches or crossing a road is a reliable way of estimating group size. The largest group verifiably observed in this way contained over 1,300 individuals, in Lopé National Park, Gabon—the largest aggregation of nonhuman primates ever recorded.[28] These groups are made of adult females and their dependent offspring.[29] Males live a solitary lifestyle, and only enter hordes when females are receptive to mating, which lasts three months each year.[21][29] All-male bachelor groups are not known to exist.[21][29]

Sleeping mother with young

The mating season of the mandrill takes place from June to October, which is when the sexual swellings of the female occur.[29] They breed every two years. When breeding, a male will follow and guard a female in estrus. Adult males exist in two different forms: the brightly colored and "fatted" dominant males, and the paler and "nonfatted" subordinate males. Both males engage in mating, but only the dominant males can sire offspring. Males sometimes fight for breeding rights which results in dominance. Though conflicts are rare, they can be deadly. Gaining dominance, that is becoming the alpha male, results in an "increased testicular volume, reddening of sexual skin on the face and genitalia, and heightened secretion of the sternal cutaneous gland".[30] When a male loses dominance or its alpha status, the reverse happens, although the blue ridges remain brightened. There is also a fall in its reproductive success. This effect is gradual and takes place over a few years.[31][32][33] When subordinates mate-guard a female, the competition between them allows the dominant males to have a greater chance of siring offspring,[34] since subordinates outnumber dominants 21 to 1. There is also a dominance hierarchy among females, with reproductive success being displayed in shorter interbirth intervals amongst these alpha figures and the beginning of reproduction at earlier ages.[34]

Mandrill births occur from January to May.[35] Most births in Gabon occur in the wet season, from January to March, and gestation usually lasts 175 days.[34] In captivity, 405 days separate each birth.[34] Young are born with a black natal coat and pink skin. The females do most of the raising of the young. Alloparenting exists in this species, with female relatives providing care for the young.[36] Males leave their natal groups when they are six years old and stay along the boundary of the social group.[21][29]

Mandrills will make a "silent, bared-teeth face", in which the teeth are bared, the head crest is erect and the head shakes. This may serve as a peaceful form of communication.[37][38] A mandrill submits by presenting its rump. With aggression, mandrills will stare, bob their heads, and slap the ground.[38] Vocalizations like roars, crowings, and "two-phase grunts" are made for long distances, while "yaks", grunts, "k-alarms", "k-sounds", screams, girneys, and grinds are made at short distances.[39]

Status and conservation[edit]

The mandrill is considered vulnerable and is affected by deforestation.[2] However, hunting for bushmeat is the more direct threat. Mandrills are particularly threatened in the Republic of the Congo.[2] Nevertheless, there have been captive-bred individuals that have been successfully reintroduced into the wild.[40]

References[edit]

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  2. ^ a b c Oates, J. F. & Butynski, T. M. (2008). Mandrillus sphinx. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 4 January 2009.
  3. ^ Linne´, Carl von (1758). Systema naturæ. Regnum animale. (10 ed.). p. 25. Retrieved 19 November 2012. 
  4. ^ a b Nowak, Ronald M (1999). Primates of the World. Baltimore, Maryland: The Johns Hopkins University Press. pp. 151–152. ISBN 0-8018-6251-5. Retrieved 7 September 2010. 
  5. ^ Darwin, Charles (1871). The Descent of Man, and selection in relation to sex. D. Appleton and Co, New York. 
  6. ^ a b c Ankel-Simons F. (2007) Primate Anatomy: an introduction, 3rd Edition. San Diego: Elsevier Acad Press.
  7. ^ Feistner, Anna T.C. (1991). "Scent Marking in Mandrills, Mandrillus sphinx". Folia Primatologica 57: 42. doi:10.1159/000156563. 
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  9. ^ Sandhyarani, Ningthoujam (30 March 2011). "Mandrill Monkey Facts". Retrieved 5 February 2012. [unreliable source?]
  10. ^ Changes in the Secondary Sexual Adornments of Male Mandrills (Mandrillus sphinx) Are Associated with Gain and Loss of Alpha Status (Joanna (Jo) Setchell) – Academia.edu. Durham.academia.edu. Retrieved on 2012-08-21.
  11. ^ Mandrill. WAZA – World Association of Zoos and Aquariums – Virtual Zoo.
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  21. ^ a b c d e f Harrison MJS (2009). "The mandrill in Gabon's rain forest-ecology, distribution and status". Oryx 22 (4): 218. doi:10.1017/S0030605300022365. 
  22. ^ a b Rogers, M. E., Abernethy, K.A., Fontaine, B., Wickings, E.J., White, L.J.T and Tutin, C.E.G. (1996). "Ten Days in the Life of a Mandrill Horde in the Lopé Reserve, Gabon". American Journal of Primatology 40 (4): 297. doi:10.1002/(SICI)1098-2345(1996)40:4<297::AID-AJP1>3.0.CO;2-T. 
  23. ^ Kudo H, Mitani M. (1985). "New record of predatory behavior by the mandrill in Cameroon". Primates 26 (2): 161. doi:10.1007/BF02382015. 
  24. ^ Tutin CEG, Ham RM, White LJT, Harrison MJS (1997). "The primate community of the Lopé Reserve, Gabon: diets, responses to fruit scarcity, and effects on biomass". American journal of primatology 42 (1): 1–24. doi:10.1002/(SICI)1098-2345(1997)42:1<1::AID-AJP1>3.0.CO;2-0. PMID 9108968. 
  25. ^ Yorzinski JL, Vehrencamp SL. "Mandrill antipredator behavior" (PDF). University of California. Retrieved 2012-08-02. 
  26. ^ Jouventin P. (1975) "Observations sur la socio-ecologie du mandrill". Terre et la Vie 29:493–532.
  27. ^ Gill, Victoria (22 July 2011). "Mandrill monkey makes 'pedicuring' tool". BBC. Retrieved 4 February 2012. 
  28. ^ "Gabon". Wildlife Conservation Society. Retrieved 4 February 2012. 
  29. ^ a b c d e Abernethy KA, White LJT, Wickings EJ (2002). "Hordes of mandrills (Mandrillus sphinx)". Journal of Zoology 258: 131. doi:10.1017/S0952836902001267. 
  30. ^ Setchell, J. M., and Dixson A.F. (2001). "Changes in the Secondary Sexual Adornments of Male Mandrills (Mandrillus sphinx) Are Associated with Gain and Loss of Alpha Status". Hormones and behavior 39 (3): 177–84. doi:10.1006/hbeh.2000.1628. PMID 11300708. 
  31. ^ "Mask of the Mandrill". PBS. November 2006. Retrieved 4 February 2012. 
  32. ^ Setchell, J. M.; Jean Wickings, E. (2005). "Dominance, status signals and coloration in male mandrills (Mandrillus sphinx)". Ethology 111: 25. doi:10.1111/j.1439-0310.2004.01054.x.  edit
  33. ^ Dixson, A. F.; Bossi, T.; Wickings, E. J. (1993). "Male dominance and genetically determined reproductive success in the mandrill (Mandrillus sphinx)". Primates 34 (4): 525. doi:10.1007/BF02382663.  edit
  34. ^ a b c d Setchell, J. M., and Dixson A.F. (2002). "Developmental Variables and Dominance Rank in Adolescent Male Mandrills (Mandrillus sphinx)". American journal of primatology 56 (1): 9–25. doi:10.1002/ajp.1060. PMID 11793410. 
  35. ^ Wickings, E. J., and Dixson, A.F. (1992). "Development from birth to sexual maturity in a semi-free-ranging colony of mandrills (Mandrillus sphinx) in Gabon". Journal of reproduction and fertility 95 (1): 129–38. doi:10.1530/jrf.0.0950129. PMID 1625228. 
  36. ^ Charpentier, M., Peignot, P., Hossaert-Mckey, M., Gimenez, O., Setchell, J.M. and Wickings, E.J. (2005). "Constraints on control: factors influencing reproductive success in male mandrills (Mandrillus sphinx)". Behavioral Ecology 16 (3): 614. doi:10.1093/beheco/ari034. 
  37. ^ Bout N, Thierry B (2005). "Peaceful meaning for the silent bared-teeth displays of mandrills". International Journal of Primatology 26 (6): 1215. doi:10.1007/s10764-005-8850-1. 
  38. ^ a b Setchell JM, Wickings EJ (2005). "Dominance, status signals and coloration in male mandrills (Mandrillus sphinx)". Ethology 111: 25. doi:10.1111/j.1439-0310.2004.01054.x. 
  39. ^ Kudo H (1987). "The study of vocal communication of wild mandrills in Cameroon in relation to their social structure". Primates 28 (3): 289. doi:10.1007/BF02381013. 
  40. ^ Peignot P, Charpentier MJE, Bout N, Bourry O, Massima U, Dosimont O, Terramorsi R, WIckings EJ (2008). "Learning from the first release project of captive-bred mandrills Mandrillus sphinx in Gabon". Oryx 42. doi:10.1017/S0030605308000136. 
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