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Mandrillus Sphinx - Sexual Dimorphism and Physical Diversity of Individuals in Relation To Social Status
Mandrillus Sphinx - Sexual Dimorphism and Physical Diversity of Individuals in Relation To Social Status
geographical range, and lifestyle. This much is immediately obvious even from distant,
secondhand study of primates. It is also paramount, however, to recognize that this variation
is not limited to species by any means, and that it extends to the individual. Social roles- in
some species more so than others- do much to dictate individual animals’ behaviors and
physical appearances. Some prominent and widespread categories of these diverse roles
stem from gender, as well as social status (particularly the dominance rank among males).
Many primate species would be effective case studies for these phenomena of intraspecific
diversity, but in this specific study, Mandrillus sphinx will be examined in detail and depth
while it is true that taxonomists once classified mandrills in the Papio genus (the genus
containing all baboons), mandrills have since been given their own genus of Mandrillus by
Groves in 1989 (“Mandrillus sphinx”). They share this genus with only one other species:
the highly endangered Mandrillus leucophaeus (common name: drill). The name “mandrill”
literally means “man-like drill,” with “drill” of course simply referring to its nearest relative,
which is an old West African name for the monkeys (“America Zoo”). Drills have had their
native home range in Cameroon and southeastern Nigeria almost completely depleted by
human development, and have consequently had their range almost entirely confined to the
island of Bioko, which belongs to the small West African nation of Equatorial Guinea. Drills
were, in fact, rated by the International Union for Conservation of Nature and Natural
Resources (IUCN) as the highest conservation priority among all African primates (Massicot
1
See (Fig. 1) for full taxonomy.
3
2006). The mandrill’s geographical range is quite broad compared to those of its more
endangered cousins, but still rather small compared to many primate species, straddling the
equator and confined roughly between the latitudes of 4°N and 4°S. Its western boundary is
the Atlantic Ocean, but the full extent of its eastern range is still ambiguous. Wild mandrills
live primarily within the borders of Gabon, Equatorial Guinea, and southern Cameroon
(south of the Sanaga River); although occasionally they have also been sighted in isolated
The mandrill is a uniquely distinctive animal. It is the largest monkey in the world3,
(Feistner 2005) usually measuring from 61-76cm in length and up to 80cm tall. Females
weigh an average of 11.5kg, but males weigh an average of 25kg. Some of the largest males
have been known to weigh up to 54kg. It is evident from this basic data that a great deal of
sexual dimorphism exists within the M. sphinx species; however, as it turns out, the
dimorphism extends beyond a difference in size. Perhaps the mandrill’s most recognizable
feature is its bright colors: this includes a red stripe down the center of the face, and light
blue ridged segments on either side of the stripe. The fur is olive-colored, the rump is bright
blue, and a yellow-colored beard hangs from the chin. These colors are far more pronounced
in mature males than in females or juveniles (Ingmarsson 1999). The physical distinctions
between sexes, as well as the distinction between males of different social rank, will be
The mandrill’s morphology allows for a diverse range of easy locomotion. Unlike
most monkeys, it has adapted to a primarily terrestrial lifestyle as opposed to an arboreal one,
although it does often shelter itself in trees at night. The combination of strong, long limbs, a
2
See (Fig. 2) for map of range.
3
This includes both Platyrrhini and Catarrhini infraorders, with “monkey” defined as any non-hominoid
anthropoid.
4
clavicle, and opposable thumbs allow it to climb trees with ease while also maintaining a
natural quadrupedal walk, which allows for their terrestrial daytime lifestyle. Their tails are
considerably shorter than those of most monkeys, which indicates less reliance on tree-
climbing4. Like most primates, they are native and best adapted to tropical rain forest
Like all Cercopithecines, mandrills are largely frugivorous and contribute to seed
dispersal in forests (Ingmarsson 1999). However, they have been known to supplement their
diet with a variety of other foods. Various researchers have observed mandrills for lengthy
periods of time and collected fecal samples to estimate the relative amounts of different types
of food the mandrill consumes, and the results of this research has varied some depending on
the location, most likely due to differing abundances of specific food resources. Specifically,
one study in the Lope Reserve in Gabon (Tutin et. al. 1997) yielded a higher percentage of
leaves (8.2%) than a study in Cameroon conducted by Hoshino (1985), while the Cameroon
study yielded a higher percentage of animals (7.6% vs. 4.1% in Gabon). Both studies,
however, revealed that mandrills have a dietary staple of about 84% fruit or fruit material5
(Flannery 2003). It is not known why mandrills consume this specific amount of fruit and
supplement the remainder of their diet with leaves, animals, fungi, and even small amounts
of soil, but it is a trend that is not uncommon among primates. Mandrills, however, are able
to eat more animals than most other monkeys due to their large size. Animals that serve as
prey for mandrills include insects, worms, snails, frogs, lizards, and on some occasions they
have even been observed to eat snakes (Ingmarsson 1999). Still, perhaps the mandrill’s most
impressive hunting feat is the predation of Cephalophus dorsalis, or duiker, a small African
4
Many monkeys use their tails for balance or grasping branches. Both functions enable a highly arboreal
lifestyle. When mandrills do need to climb trees, they utilize their opposable thumbs.
5
Includes seeds and pith.
5
antelope species (Kudo & Mitani, 1985). Predation of such large vertebrates is not common
and has only been observed on isolated occasions. Another unique dietary trait of the
mandrill is that, due to its terrestrial lifestyle, it mainly eats fruit that has fallen off the tree
already. Some have speculated that this accounts for why the mandrill has a large home
range when compared to other primate species in similar environments (Hoshino et al.,
1984). It is not entirely known whether mandrills have any significant predators, but some
evidence indicates that Panthera pardus (leopards) might prey on mandrills. This evidence
comes from analysis of leopard feces, although it remains possible that the leopard was
The mandrill, though not technically endangered like its previously discussed relative,
the drill, is classified as “vulnerable,” which means that it is at risk of becoming endangered.
In recent years the mandrill has suffered a population decline, primarily due to habitat
destruction by humans. A more minor, but not insignificant problem that humans have
caused for the mandrill is from hunting. Mandrills are sometimes locally hunted for food.
Because of their loud vocalizations, they often make themselves conspicuous to these
hunters. Additionally, mandrills have a reputation as pests to some West African farmers. As
a result of too much habitat destruction, mandrills are occasionally forced to “steal” crops.
This unfavorable reputation may contribute to a harmful lack of effort to preserve the species
(Ingmarsson 1999).
organization of mandrills is a harem structure, in which there is a single large, dominant male
who is polygynous. The male has exclusive mating rights over the females who make up the
remainder of the group, and these groups have been known to contain up to 15 individuals.
6
However, sometimes mandrills are found in extremely large groups, ranging up to several
hundred individuals. These groups consist of many males and many females. Breeding is
infrequent, occurring only about once every two years, and females typically give birth to
only one offspring at a time. Mating season is in the summer, and the gestation period lasts
The intricacies of the mandrill social structure will be under close examination for the
remainder of this paper. Specifically, the process under which a male may rise to dominance
will be investigated, as will the events that occur to those males that never do become harem
leaders. The sexually dimorphic nature of mandrills, which are believed to have more sexual
dimorphism than any other primate species (Charpentier et al. 2004), is not to be overlooked
when investigating these subjects and it in fact will be another main focal point of this case
study. The relationship, if any, between the male dominance in this species and the extreme
Juvenile males have yet to grow to the comparatively huge size of their adult
counterparts, nor have they yet acquired the colors. Therefore, it is apparent why the
adolescent life stage for males would be of great interest. Charpentier et. al. explore this
realm in a 2004 experiment involving the social interactions of eleven adolescent males. As
the researchers mention, mandrills are a good choice for an experiment such as this because
the adolescent phase is very long, lasting from approximately four to ten years of age.
During this time, male mandrills undergo major growth spurts, and the colors on the face,
rump, and genitalia all brighten. Charpentier et al. observed that among the 11 adolescents,
the eight younger individuals formed a cohort, while the other three were more solitary and
more aggressive. Female adolescence lacks this long, eventful phase, presumably because
7
they do not require nearly as much change from juvenile to adult. Female adolescents are in
fact difficult to define in mandrills because so little change occurs, save for the obvious
development of gametes. Due to this difficulty, the category of “female adolescents” was
The experiment categorized types of behavior, which was recorded through close
observation of all eleven individuals, scored against control groups involving every other
results showed that the rise to male dominance was as gradual and defined as physical
development might indicate. One key lesson of the experiment is that “adolescent” is
perhaps not a sufficient categorization for a mandrill between the ages of four and ten. The
researchers found that two distinct levels of male mandrill adolescence exist. The first,
exemplified by the cohort of eight younger individuals, begins around the age of four, when
the testicles descend. This life stage is categorized by mainly conspecific interaction among
others in the same age group, including a heavy emphasis on playing. Once the mandrills
age past that stage, however, they begin to engage in primarily aggressive interaction towards
virtually every other group, with the notable exception, unsurprisingly, of adult males. This
was exemplified by the three older adolescents not included in the previous cohort.
Adult males, however, as seen in Charpentier et al. (2004, p. 70), exert by far the
smallest amount of aggression on other group members despite receiving the bulk of it.
Perhaps it is safe to assume that this is due to his duty as the protector of the group, or harem,
if applicable. This leads to the realization that because of their unrivaled combination of
strength and aggression, older male adolescents are by far the most dangerous members of
the group, and in general the most dangerous mandrills toward other mandrills. It is
8
interesting to speculate on the reasons for this, but one possibility is that it serves as a tactic
to gain acceptance. Male adolescents vie for dominance, and gain the respect of fellow
group members through aggressive behavior. The issue of dominance among wild mandrill
groups will be discussed further in subsequent studies, which will be carefully analyzed.
Although Charpentier et al.’s study brought to light some informative data about the
details of male mandrills’ growth and development, one factor in their study was largely
ignored- social rank. Setchell and Dixson (2002) also observe the male mandrill’s growth in
the adolescent phase, but they concentrate on the mandrills’ relative social status. It was
observed that the males that rose to dominance and were more social had larger testes and
higher testosterone levels, while the males that were more “peripheral and solitary” (Setchell
& Dixson 2002, p. 9) had smaller testes and lower testosterone levels. The study measured
13 males aged five to nine years at the International Medical Research Center in Gabon over
a 21-month period (Setchell & Dixson 2002, p. 10), and measured various characteristics in
these males and the degree to which they changed over time. These traits included, among
others, body mass, testosterone levels, the brightness of the male sexual colors on various
parts of the body, and the age at which they left their natal groups. Setchell and Dixson
ultimately concluded that “Marked individual differences in the timing of testicular function,
sexual skin coloration, and other features are also apparent in males, as well as differences in
the ages at which they emigrate from their natal groups. These differences correlate most
strongly with male dominance rank, so that high-ranking males develop faster and emigrate
earlier than subordinate males in mandrill social groups (Setchell & Dixson 2002, p. 23).
This data may seem unsurprising, considering that the traits most unique to males
were more pronounced in the males that rose to dominance in the social structure. It seems
9
clear that there is a genetic component to the factors that determine male dominance.
However, Charpentier et al.’s study, discussed earlier, revealed that essentially all adolescent
males go through a solitary and sometimes aggressive phase, usually between their sixth and
tenth years of age. Since this phase is virtually universal among male mandrills, it would
seem to suggest that all male mandrills make an attempt to rise to dominance. The purpose
of the solitary phase in adolescents is thought to be for males to develop better survival skills,
which they would need in adulthood especially if they were to become dominant males over
a harem. As mentioned earlier, a harem structure consists of one dominant male and up to
fifteen females with whom he has exclusive mating rights. Common sense regarding simple
gender ratios should tell us that it is not possible for all males to rise to this level of
competition is present among males approaching adulthood, mostly over mating prospects.
In summation, it appears likely that male dominance is determined by both genetics and by
competition.
Two reasonable questions that may now be asked are simply; why do mandrills have
this male-dominant harem structure, and why do they exhibit such extreme sexual
dimorphism? It is easy to answer these questions one at a time with the assumption that the
other condition is already in place. For instance, it is not difficult to imagine why males
having a much greater body size than females might make them more dominant, and it is
easy to see why a harem structure works better when this natural sign of dominance exists.
However, the question becomes much more difficult if neither sexual dimorphism nor the
harem social structure is assumed to have previously existed. What might cause these
Firstly, it is important that mandrills’ sexual dimorphism with regard to body size is
not unique among Cercopithecines. For examples, males may be up to twice the size of
females in Papio hamadryas, and the males also have a large, bushy mane that the females
lack (“Hamadryas baboon, sacred baboon” (BBC)). In fact, all baboons are dimorphic in
size. Mandrills were thought for a long time to be baboons (members of the Papio genus),
but were separated from them recently. Nonetheless, they continue to share many
characteristics with them. It turns out that Papio hamadryas baboons also share the
mandrill’s harem social structure. Now it is evident that the concurrence of sexual
dimorphism and male-dominated social structures may be a trend among some monkeys.
Perhaps a logical reason for these phenomena concurring in many Cercopithecines lies within
their life histories. It was previously mentioned that mandrill females typically give birth to
only one offspring at a time, yet their gestation periods are fairly long and females only mate
approximately every two years. This may be a legitimate reason for the harem structure.
Normally, such statistics would be foretelling of a low birth rate, which could lead to
population declines6. Harem structures, however, by making a single male readily available
to all females in mating season, would allow females to reproduce as often as they are ready.
This explanation for harem structures in mandrills would make it clear why sexual
dimorphism in body size is necessary. It ensures male dominance within these harem
structures and allows a maximum number of total offspring to be produced for group. It also
eliminates the element of competition for mates within groups, although males may still
possibly compete for social status during their adolescent years, as mentioned before. Even
6
It was previously mentioned that mandrill population is suffering a decline. This is due mostly to human
activities including hunting and land development. Mandrills could be especially susceptible to this because
they have very few natural predators and may be unaccustomed to hiding or defending themselves.
11
under this hypothesis, ambiguities abound. For instance, Setchell et al. (2006) recently
observed an event that raises some questions about the degree to which male dominance is
absolute. Setchell et al. chronicle the observation of several female mandrills attacking a
dominant male who had been injured by another male. Also mentioned in their report was
that in previous observations, large groups of female individuals had even been reported to
kill a male (Setchell et al. 2006, p. 411). The researchers concluded that this observation
“suggests that females have a degree of control over male group membership, despite the
large degree of sexual dimorphism in this species” (Setchell et al. 2006, p. 411). These
isolated incidents are puzzling given what we know about sexual dimorphism’s contribution
arguably the mandrill’s most defining feature- the bright colors that appear on the male
during puberty.
The presence of these colors is what most notably distinguishes the mandrill from its
closest relative, Mandrillus leucophaeus, or the drill. As reported earlier, the brightness of
these colors correlates to dominance. Darwin, in fact, used the mandrill as an example for
his theory of sexual selection, and studies have indicated that females do indeed usually
prefer brightly colored males. If the mandrill and the drill diverged from a common ancestor,
it is reasonable to ask what prompted the development of the mandrill’s bright colors, and
whether they have anything to do with the mandrill’s superior success at survival than drills,
which are among the world’s most endangered primate species. Andreas Paul (2001)
explored this unanswered question of evolution. Although no conclusion has been reached,
Paul offers some theories as to how the change may have come about. Paul states that
Darwin’s theory that females prefer brightly colored males is “unproven,” but states an
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alternative explanation made by Wickler in 1967, in which Wickler hypothesized that the
function of the colors was instead to have a “’demoralizing effect’ on possible rivals”
If Wickler’s hypothesis is accurate, brightly colored males would still have a greater
chance of mating because they would have the advantage in competition with other males-
not because of sexual selection. If Darwin’s theory of sexual selection applies to the
mandrill, then evolution is seemingly more complicated, because not only would males have
to evolve the colors, but females would have to evolve a preference for the colors. This
seems illogical if the colors did not yet widely exist around the time when mandrills diverged
from drills. Wickler’s hypothesis seems more likely, only because it requires a single
evolutionary event in males which the females would subsequently evolve to adapt to, as
opposed to Darwin’s theory in which male and female change would be mutually dependent
upon one another. Some researchers including Setchell and Dixson have documented an
apparent higher likelihood of reproductive success for brightly colored males, but to
determine the cause of this success would be far more difficult. Perhaps someday an
experiment could be arranged to determine the nature of this remarkable phenomenon unique
to Mandrillus sphinx.
Literature cited
Charpentier, M., Peignot, P., Hossaert-McKey, M., and Wickings, E.J. 2004. Changes in
Social Interactions During Adolescence in Male Mandrills (Mandrillus sphinx). American
Journal of Primatology. Vol. 63(2), 63-73.
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Feistner, Dr. Anna. April 02 2005. “Mandrill- Mandrillus sphinx” (On-line), ARKive.
Accessed November 06, 2006 at
http://www.arkive.org/species/GES/mammals/Mandrillus_sphinx/.
Flannery, Sean. October 09, 2003. “Mandrill- Mandrillus sphinx” (On-line). Accessed
November 06, 2006 at http://members.tripod.com/uakari/mandrillus_sphinx.html.
Kudo, Hiroko, and Matani, Masazumi 1985. New record of predatory behavior by the
mandrill in Cameroon. Primates. Vol. 26(2), 161-167.
Massicot, Paul. August 01 2006. “Animal Info- Drill” (On-line), Animal Info. Accessed
November 01, 2006 at http://www.animalinfo.org/species/primate/papileuc.htm.
Paul, Andreas. 2002. Sexual Selection and Mate Choice. International Journal of
Primatology. Vol. 23(4), 877-904.
Setchell, Joanna M., and Dixson, Alan F. 2002. Developmental Variables and Dominance
Rank in Adolescent Male Mandrills (Mandrillus sphinx). American Journal of Primatology.
Vol. 56(1), 9-25.
Setchell, Joanna M., Knapp, Leslie A., Wickings, E. Jean. 2006. Violent coalitionary attack
by female mandrills against an injured alpha male. American Journal of Primatology. Vol.
68(4), 411-418,
Tutin, C.E.G., Ham, R.M., White, L.J.T., and Harrison, M.J.S. 1997. The primate communtiy
of the Lope Reserve, Gabon: Diets, responses to fruit scarcity, and effects on biomass.
American Journal of Primatology. Vol. 42, 1-24.
Wickler, W. 1967. Socio-sexual signals and their intra-specific imitation among primates. In
Morris, D. (ed.), Primate Ethology,Weidenfield and Nicholson, London, pp. 69–147.
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Order Primates
Suborder Anthropoidea
Infraorder Catarrhini
Superfamily Cercopithecoidea
Family Cercopithecidae
Subfamily Cercopithecinae
Genus Mandrillus
Species sphinx