Chlorocebus sp.
AT A GLANCE
Status: Least concern (all other sp.), Data deficient (Ch. djamdjamensis)
Life span: 11 to 13 years (captive)
Total population: Unknown
Regions: Sub-Saharan Africa
Gestation: 5.5 months (165 days)
Height: 490 mm (M), 426 mm (F)
Weight: 5.5 kg (M), 4.1 kg (F)
TAXONOMY
Suborder: Haplorrhini
Infraorder: Simiiformes
Superfamily: Cercopithecoidea
Family: Cercopithecidae
Subfamily: Cercopithecinae
Genus: Chlorocebus
Species: aethiops, cynosuros, djamdjamensis, pygerythrus, sabaeus, tantalus
Subspecies: Ch. p. excubitor, Ch. p. hilgerti, Ch. p. nesiotes, Ch. p. pygerythrus, Ch. p. rufoviridis, Ch. t. budgetti, Ch. t. marrensis, Ch. t. tantalus
Other names: Ch. aethiops: Cercopithecus
aethiops, Cercopithecus
aethiops aethiops, or Chlorocebus aethiops aethiops; grivet
or savanna monkey; singe vert (French); grünmeerkatze (German); mono verde (Spanish); grön markatta
or vervett (Swedish); Ch. cynosuros: malbrouck; Ch.
djamdjamensis:
Bale Mountains vervet or djam-djam; Ch. pygerythrus: Cercopithecus
aethiops pygerythrus or Chlorocebus aethiops pygerythrus; vervet
monkey; Ch. sabaeus: Cercopithecus aethiops sabaeus or Chlorocebus
aethiops sabaeus; green monkey; Ch. tantalus: Cercopithecus
aethiops tantalus or Chlorocebus aethiops tantalus; tantalus monkey
The classification of vervet monkeys was recently updated, moving all of the
species from the genus Cercopithecus to a new genus, Chlorocebus (Rowe
1996; Groves 2001). There are now at least six species of vervets recognized,
but often in the literature, they are ubiquitously referred to as Chlorocebus
aethiops (Ch. aethiops) by their former genus, Cercopithecus
aethiops (C. aethiops) (e.g. Grubb et al. 2003).
They are sometimes lumped together with a group of primates called guenons,
medium-sized arboreal
African monkeys of the genus Cercopithecus (Lernould
1988; Oates 1996). Groves (2001) recommends further revision of this genus
and in the future, there will likely be more species and subspecies identified
(Grubb et al. 2003).
MORPHOLOGY
Typically, Ch. aethiops, Ch. pygerythrus, Ch. tantalus
and Ch. cynosuros have
a yellow to greenish-brown coat with white undersides and white fur on their
brows and cheeks. They have bluish skin on their abdomens while their faces,
hands and feet are all black-skinned (Rowe 1996; Groves 2001). Males of all
species of vervets have bright blue scrotal areas contrasting with red penises.
This striking coloration, when combined with the white fur of their undersides
is highlighted in what is referred to as the "red, white, and blue display"
(Fedigan & amp; Fedigan 1988; Rowe 1996). There is some variation in fur length
and coloration among the species. Chlorocebus sabaeus has golden-green
fur with pale hands and feet. The tip of the tail is golden yellow as are the
backs of the thighs and cheek whiskers. They do not have a distinguishing band
of fur on the brow, like other species, and males have a pale blue scrotum (Groves
2001). Bale Mountains vervets, Ch. djamdjamensis, have much
longer, thicker fur than other species and their fur is darker brown. They have
dark gray hands and feet and white beards and there is only a faint white brow
band in this species (Groves 2001). Infant vervets are born with black
natal
coats and pink faces that gradually change to the adult coloration by 12 weeks
of age (Lee 1984).
Chlorocebus pygerythrus
All vervet males and females are sexually dimorphic and wild adult males weigh
between 3.9 and 8.0 kg (8.60 and 17.6 lb), averaging 5.5 kg (12.1 lb), and measure
between 420 and 600 mm (1.37 and 1.97 ft), averaging 490 mm (1.61 ft) from the top of the head to the base of the tail. Wild adult females weigh between
3.4 and 5.3 kg (7.50 and 11.7 lb) and average 4.1 kg (9.04 lb), and measure between 300 and 495 mm (11.8 in and 1.62 ft), averaging 426 mm (1.40 ft) (Napier 1981; Skinner & Smithers 1990).
Vervets move quadrupedally
and they are equally as comfortable on the ground
as they are in the trees. They are
semi-terrestrial
and semi-arboreal, spending
time feeding and traveling on the ground during the day and retreating to the
trees to sleep at night (Fedigan & amp; Fedigan 1988). They have the characteristic
cheek-pouches like other members of the superfamily Cercopithecoidea which
allow them to forage and store food to be eaten later (Rowe 1996).
Lifespan in wild vervets is difficult to characterize because of high rates
of predation in the long-term study areas. One female was at least 13 years
old when she died and females in captivity have lived this long. Between 11
and 12 years, serious health problems are seen in captive vervets and this
is probably the upper limit of their lifespan (Fairbanks & amp; McGuire 1986).
RANGE
Vervets are the most widespread of the African monkeys and inhabit large parts
of sub-Saharan Africa (Wolfheim 1983; Lernould 1988). They are found broadly
across the continent from Senegal to Ethiopia and north into Egypt and Eritrea
as well as southward to South Africa. Vervets are largely absent from the forests
of the Congo Basin in west-central Africa, though some species inhabit the
edges of these forests (Wolfheim 1983). The species are separated geographically
but some areas of hybridization occur
(Groves 2001). Political instability in some regions in eastern and western Africa
has made censusing vervets
difficult, but the range is reasonably defined (Zinner et al. 2002). Chlorocebus
pygerythrus ranges
from the Ethiopian Rift Valley in central Ethiopia eastward into Somalia, and
southward into Kenya, northern Tanzania and eastern Uganda. To the north, Ch.
aethiops is
found in Sudan, east of the White Nile River, Ethiopia, Eritrea, and probably
into southeastern Egypt. In the southeastern part of its range, Ch. aethiops hybridizes
with Ch. pygerythrus as well as with Ch. p. hilgerti at the Omo River
in Ethiopia (Isbell & amp; Enstam under review). Another species of vervet found
in Ethiopia is Ch. djamdjamensis, restricted to the Bale Mountains
region and surrounding highland areas east of the central Rift Valley. Chlorocebus
tantalus is found in Sudan, Uganda, and northwestern Kenya around Lake
Turkana, one of
Africa's
Great Lakes,
and its range stretches west into Togo, Benin, Nigeria, Niger, Chad, Cameroon,
Equatorial Guinea, Central African Republic, Congo, and into Ghana where it is
restricted by the Volta River. It hybridizes with Ch. p. rufoviridis in Uganda
along the northern and western shores of Lake Victoria (Isbell & amp; Enstam under review). The westernmost species of vervet is Ch. sabaeus,
found from Senegal to the west bank of the Volta River in Ghana and ranging in
Mauritania, Mali, Gambia, Guinea-Bissau, Guinea, Sierra Leone, Liberia, Côte
d'Ivoire, and Burkina Faso. Chlorocebus cynosuros is found in northern
Namibia, Angola, southern Democratic Republic of Congo, Botswana, and in
Zambia (Groves 2001). Chlorocebus is also found in Rwanda, Burundi,
South Africa, Lesotho, Zimbabwe, Swaziland, Malawi, and Mozambique (Seier
2003).
Chlorocebus sp. range (in red)
In addition to being found in 39 African nations, one species of vervet is
also found on the Cape Verde islands off the horn of Africa in the Atlantic
Ocean and on several Caribbean islands in the West Indies. Chlorocebus
sabaeus was introduced in the late 1600s to the islands of St. Kitts,
Nevis, and Barbados when ships involved in the slave trade traveled from to
the Caribbean from West Africa (van der Kuyl et al. 1996). Green monkeys traveled
as pets or as items to be traded and sold and have since adapted well to life
on these islands where they are sometimes considered a pest species (Fedigan & amp; Fedigan
1988; Boulton et al. 1996).
Dorothy Cheney and Robert Seyfarth have conducted long-term research on vervets
at Amboseli National Park in southern Kenya since 1977. Lynne Isbell has also
been conducting long-term research on vervets at Amboseli and with Karin Enstam at a private
conservation area, Segera Ranch, in north-central Kenya since 1992. Vervets
have also been studied in the Caribbean for several decades. Most of the current
published information about vervets is from studies done at these sites.
HABITAT
Vervets are habitat generalists, as is obvious from their widespread range in
Africa and the success of introduced populations. They are tolerant of a wide
variety of habitats and can live in humid rainforests, semi-desert environments,
or swamps from sea level to elevations up to 4500 m (14,764 ft); their only
limitation seems to be water availability and the presence of sleeping trees
(Wolfheim 1983; Chapman & amp; Fedigan 1984; Fedigan & amp; Fedigan 1988). Because
of this limitation, they are especially prevalent in riverine forests bordering
savannas. They are also able to exploit areas near cultivated fields because
they are adept at raiding crops (Struhsaker 1967; Oates 1996; Isbell et al.
1998). Additionally, vervets can survive quite well in urban areas (Wolfheim
1983; Shimada & amp; Shotake 1997). Vervets are rarely found in the depths of
dense forests, but rather utilize the edges of tropical rain forests, lowland
evergreen forests and montane forests. They seem to prefer wooded rather than
heavily forested areas, such as dry deciduous forest, scrub forests and gallery
forests, which are composed of both trees and shrubs (Nakagawa 1999). They
are not found in open grassland with no trees, but they spend some time in open
savannas moving between wooded areas (Wolfheim 1983).
Data on climate have been provided for study sites within their range. In
Senegal, there are two seasons; the dry season lasts from November to May
and the rainy season lasts from June to October. The mean annual rainfall is
954 mm (3.14 ft), and almost all of it falls in the months from June to October.
The dry season has maximum temperatures between 33º and 40º C (91.4º and
104º F) while the rainy season is slightly cooler, with average maximum
temperatures ranging from 30º to 33º C (86º to 91.4º F)
(Harrison 1984). In Cameroon, the wet season lasts from May to September
while the rest of the year is dry. The average annual rainfall is 497 mm (1.63
ft) and average monthly temperatures range from 22.7º to 33.7º C
(72.9º to 92.7º F) (Nakagawa 1999). The climatological conditions
in which vervets live in Kenya include a drier season lasting from September
to January or March with average annual rainfall reaching 700 mm (2.30 ft)
(Isbell et al. 1999).
Chlorocebus sabaeus
In the West Indies, vervets inhabit both densely populated areas in urban
habitats as well as highly cultivated agricultural habitats with lower human
population density (Hoorocks 1986). Because agriculture and fuelwood extraction
have significantly decreased in the last 30 years, vervets also inhabit thickly
wooded ravines of secondary forest (Boulton et al. 1996). They can be found
in mangrove swamps, in stands of sea grape, a tropical evergreen, and in pasturelands
bordering wooded areas (Chapman & amp; Fedigan 1984). The rainy season lasts
from July to September and the dry season stretches from December to May. The
average temperature on St. Kitts is 23.9º C (79º F) (Poirier 1972;
Hoorocks 1986).
ECOLOGY
In much of their range, seasonal availability of food and water necessitates
the ability of vervets to forage for and consume a wide variety of foods. In
fact, vervets, along with chimpanzees (Pan troglodytes
and P. paniscus)
and baboons (Papio species),
have the most omnivorous diet of all primates. Vervets eat all of the types
of food that any primate is known to eat including leaves, gum, seeds, nuts,
grasses, fungi, fruit, berries, flowers, buds, shoots, invertebrates, bird eggs,
birds, lizards, rodents, and other vertebrate prey. They have a strong preference
for fruit and flowers, which are seasonal resources, and from month to month
vervets vary their diet tremendously to cope with fluctuations in food availability
(Harrison 1984; Fedigan & amp; Fedigan 1988). Brushfires during the dry season
often temporarily reduce vegetative cover so that little is available during
the beginning and middle of the rainy season (Adeyemo 1997). Subsequently, vervets
can suffer severe nutritional stress accompanied by hair loss and changes in
skin color (Isbell 1995). In more developed and agricultural areas, vervets
are less subject to nutritional stress and seasonal availability of food because
they depend on sources of food provided by humans including cultivated fruits,
vegetables, and cereal crops (Saj et al. 2001). On Barbados, vervets consume
fruits and vegetables grown in subsistence gardens and on larger plantations.
Some of the most damaged crops include corn, sweet potato, bananas, mangos,
papayas, guavas, cherries, cucumbers, peanuts and yams (Saj et al. 2001; Boulton
et al. 1996). They have become an increasing nuisance to farmers that grow these
crops and one management option has been to trap crop-raiding vervets and sell
them research facilities for use in biomedical research (Horrocks & amp; Baulu
1994).
In addition to varying their diet according to environmental conditions, vervets
change their daily activity patterns depending on the season. Generally, they
travel, feed, and sleep as a group (Harrison 1983). In the rainy season, vervets
spend their mornings, from about 7:00 a.m. to 11:00 a.m., traveling, feeding
and drinking water and the afternoons, from 11:00 a.m. to 3:00 p.m., are spent
feeding, drinking water, resting, grooming and doing some traveling (Adeyemo
1997). The proportion of time spent doing each activity changes during the dry
season so that less time is spent traveling and feeding and more time is spent
drinking, resting, and grooming in the mornings. During afternoons in the dry
season, more time is spent drinking water, feeding and grooming than during
the rainy season. The evenings, from 3:00 p.m. until 7:00 p.m., are spent feeding
and traveling to the sleeping tree (Adeyemo 1997). Sleeping trees utilized by
vervets average 7.7 m (25.3 ft) in height and are usually found in wooded areas
rather than standing alone or in small clusters of trees amidst open grassland.
One reason vervets use sleeping trees is to decrease the risk of predation (Nakagawa
1999). In their natural habitat, potential predators include lions, leopards,
cheetahs, other felid predators such as African wild cats, servals and caracals,
hyenas, black-backed jackals, raptors
and baboons (Papio species). In the West Indies, dogs
are serious predators and in most areas throughout their range, humans hunt
vervets either for meat or as a means to control the population (Isbell & amp; Enstam
2002; Zinner et al. 2002).
Chlorocebus pygerythrus
Like changes in their diet and activity pattern, the day range length and home
range size of vervet groups changes seasonally as well. In Senegal, vervets
travel between 665 and 2670 m (.413 and 1.66 mi) during a single day (Harrison
1983). Home range size and population density varies based on the spread of
resources available in each habitat. The smallest home ranges are seen among
vervets on St. Kitts and average .06 km2 (.023 mi2) and the largest are seen
among vervets living in Senegal, where home ranges are as large as 1.78 km2
(.687 mi2) (Harrison 1983). Where there are few resources, like in Senegal, vervets need to range farther to survive and fewer animals are supported by the ecosystem. On St. Kitts, where agricultural foods are readily available and in condensed areas such as on plantations, large numbers of animals can be supported in a much smaller space (Harrison 1983). Population densities range from 9 individuals per square kilometer (5.6 per square mile) at the Segera Ranch in Kenya to 255 individuals per
square kilometer (158 per square mile) on St. Kitts (Harrison 1983; Isbell & amp; Enstam under review). When groups encounter
one another at the border or overlapping areas of their home ranges, interactions
can be described as friendly, where the two groups temporarily merge, or as
agonistic, with serious fighting between members of both groups (Cheney 1981;
Chapman & amp; Fedigan 1984). The expression and degree of territoriality among
vervets is facultative and conditions such as habitat type, season, intensity
of predation, distribution, abundance, and defendability of resources, and the
history of relationships between the groups are all factors contributing to
the nature of the interaction of two groups (Kavanaugh 1981; Chapman & amp; Fedigan
1984). In environments with clustered, easily defendable resources, territoriality
is economically sound, especially in highly seasonal environments; vervets have
much to lose by allowing neighboring groups into their range to exploit their
food resources, especially at certain times of year. In other types of environments,
such as savanna woodlands, resources are widely distributed and would be difficult
for one group to defend without serious energetic costs, therefore, territorial
behavior is not expected under these conditions (Kavanaugh 1981).
SPECIAL NOTES
Vervets are among the most used primates in biomedical research in
the United States and abroad (Carlsson et al. 2004). Because they are
small, easily handled, nonendangered, evolutionarily closely related to
humans, and easily bred in captivity, vervets are a popular species for
use in biomedical primate research (Ervin & Palmour 2003).
Specifically, vervets are important in studying high blood pressure and
AIDS. They are one of the few species of nonhuman primates that
naturally develops high blood pressure and simian immunodeficiency virus
(SIV), the ancestor of human immunodeficiency virus (HIV), is widespread
throughout wild populations (Chakrabarti 2002; Ervin & Palmour 2003).
There is compelling evidence that SIV was transferred from monkeys or
apes in Africa to humans and led to the emergence of HIV/AIDS in humans.
Studying naturally occurring SIV and the origins of HIV/AIDS in vervets
and other African primates may help scientists discover a cure or
vaccine for the disease (Chakrabarti 2002).
Content last modified: January 3, 2006
Written by Kristina Cawthon Lang. Reviewed by Karin Enstam.
Cite this page as:
Cawthon Lang KA. 2006 January 3. Primate Factsheets: Vervet (Chlorocebus) Taxonomy, Morphology, & Ecology. <http://pin.primate.wisc.edu/factsheets/entry/vervet/taxon>. Accessed 2008 May 16.