Describer (Date): A. Sparrman (1779). "A reference to the African
buffalo." Kongl. Svenska Vet.-Akad. Handl. (Stockholm), 40:79.
Kingdom: Animalia Phylum: Chordata Class: Mammalia Order: Artiodactyla* (Even-toed hoofed animals: includes pigs, sheep goats, cattle, deer) Suborder:Ruminantia Family:
Bovinae Genus:Syncerus Species:S. caffer Subspecies:S. caffer caffer (Cape, or savanna, buffalo) S.
caffer nanus (forest buffalo)
*New anatomical and DNA evidence on the relationship between Artiodactyla (even-toed ungulates) and Cetacea (whales and dolphins) recently led to a merging of the two orders into a new group, Cetartiodactyla (Montgelard, 1997; reviewed in Kulemzina, 2009). As of October 2012, experts had not agreed on whether to define Cetartiodactyla as an official taxonomic order that would replace Artiodactyla and Cetacea. Some continue to list the forest buffalo in the order Artiodactyla (Franklin, 2011) or use the term Cetartiodactyla without defining it as an order (IUCN, 2008).
Taxonomy and Phylogeny
The tribe Bovini, which includes cattle, bison, and buffalo, seems to
have a Eurasian origin.
Closest living relative of the African buffalo on morphological and
behavioral grounds is the Asian water buffalo, Bubalus bubalus.
Syncerus and Bubalus are also closely linked by
ribosomal-DNA data (Wall et al.,
Nearest known relative of the Syncerus lineage is the extinct Ugandax
gautieri, first found in Uganda. Syncerus probably
originated in the area between Lake Victoria and the Horn of Africa,
spreading southward and westward.
Large savanna buffalo may represent recent expansion and evolution of
the smaller forest buffalo. Larger body and horn size of the savanna
buffalo may have evolved because of increased male competition in
larger herds that could be supported by range that is more open.
Nutritional superiority of a grassland diet over a forest diet may
have also been a factor in the development of larger body size.
Controversy surrounding the relationship of the various types of
African buffalo is exemplified by the 43 specific and subspecific
names that Allen (1939) listed as having been assigned to them. In
addition to S. c. caffer and S. c. nanus, other possible
subspecies include aequinoctialis, brachyceros, mathewsi,
and planiceros. East (1998) argues that aequinoctialis,
brachyceros, caffer,and nanus should be
distinguished for purposes such as assessment of conservation status.
Syncerus caffer shows a great deal of morphological variation,
and some taxonomists have regarded the forest buffalo as a separate
species from the Cape buffalo. However, there is considerable
intergradation and hybridization of the two in many areas where their
distributions overlap, and most authorities consider both to be
subspecies of S. caffer.
Syncerus originates from the Greek sun (together) and
keras (horn); a reference to the horns which are close together at
the base). Syncerus was coined by Hodgson in 1847.
caffer is derived from the new Latin cafer (of Caffraria,
or Kaffraria) and is attributed to Sparrman (1779).
Buffalo originates from the Portuguese name for the species, bufalo.
Native names for African buffalo include: the Kiswahili nyati
and mbogo; imboogo (Luhya); mboho, mbowo,
and boo (Kichagga); nyahi (Kirabai); njari
(Kuamba); jowi and jubi (Lwo); odru (Lugbara and
Madi); soyet (Kipsigi); soet (Kalenjin); ekosobwan
(Ateso); losowan (Samburu); olosowaan and olarro
(Masai), gardas (Kiliangulu), and the Somali gessi.
Bovids (including antelopes, cattle, goats, sheep, and
their relatives) are native to Africa, Asia, Europe,
and North America (Nowak,
Historical range of the African buffalo comprised nearly
all of sub-Saharan Africa, except the more arid eastern
and southwestern areas
Buffalo do not occur more than
about 20 km from water; thus, populations are often
dispersed and localized
Distribution of the African buffalo has been reduced and
fragmented by diseases (notably rinderpest), hunting,
and habitat loss (particularly agricultural development
and competition from domestic cattle)
The species has
largely disappeared from South Africa and no longer
occurs in much of western and central Africa
Reintroduced to Bontebok and Golden Gate Highlands
national parks in South Africa
Require abundant grass, shade, and water
Prefer a mosaic of
savannas interspersed with patches of forest, reeds, or scrub
Occasionally use open grassland during their seasonal movements
Found on mountain slopes up to 4,800 m above sea level
Home ranges vary:
2-4 sq km for old bulls
1,075 sq km for a large
Seasonally, herds may slightly overlap and vary in size to
maintain access to food and water, but strong attachments to
traditional areas are kept throughout the year, even during poor range
Adult Body Weight: 265-750 kg (females) to 270-870 kg (males). The Cape
buffalo S. c. caffer is the largest African bovid, 350-870 kg;
whereas, the forest buffalo S. c. nanus is smaller, weighing
265-320 kg. Body size seems positively correlated with abundance of
grasses in a particular habitat.
Adult Body Length: 215-249 cm (females) to 225-267 cm (males) in Cape
buffalo; ca. 220 cm in forest buffalo (gender not specified in the
literature reviewed for this account).
Adult Shoulder Height: 123-150 cm (females) to 125-158 cm (males). Cape
buffalo generally stand 135-170 cm in height, while forest buffalo
usually stand 97-120 cm.
Horns of females are thinner and usually shorter; males develop a
massive shield (boss) across the forehead. Horns measure 30-75 cm in
forest buffalo and 50-150 cm in Cape buffalo; larger horns are more
prone to becoming tangled in forest vegetation.
Coloration includes various shades of black, brown, red, and yellow;
the underside and chin are often pale. Herds often contain individuals
of different coloration. Color tends to darken with age, and females
do not attain the same degree of blackness as males. Darker color is
thought to be more conspicuous than lighter color, and cryptic
coloration in a large, social species may be of little value. Forest
buffalo are typically lighter (red-brown to brown) than Cape buffalo,
though forest buffalo males are sometimes black.
Pelage of juveniles is thicker than that of adults; dark brown hide
shows through on old bulls.
Forest buffalo usually have a short dorsal mane.
Other Physical Characteristics
African buffalo and Asian water buffalo have horns that are
triangular-shaped in cross-section (as opposed to round-shaped in
cattle and bison). In the African buffalo's skull, the vomer and
palatine bones remain separate, while in the water buffalo they are
Stout legs to support its ox-like body. Front hooves are larger than
the rear hooves, which reflects the weight it carries in its massive
head, horns, and neck. Broad hooves, well-developed false hooves, and
lack of an interdigital cleft are advantageous on wet ground.
Capable of running at speeds up to 48-56 km/hour, enough to outrun
lions, but not adapted to running far.
Skin may be >1 cm thick.
Females have two pairs of inguinal mammae.
No scent glands.
Loss of body water through evaporation can be reduced by allowing the
body temperature to gradually rise through the day; the "stored
heat" is then lost at night.
Stiff, relatively-immobile lips prevents it from feeding selectively on
short grass as do smaller herbivores, but a wide incisor row, along
with use of the tongue, allow it to pluck whole bundles of grass and
thus consume more quickly.
Dental formula is: incisors, 0/3; canines, 0/1; premolars, 3/3; molars,
3/3; total, 32.
Olfaction is good; hearing and particularly vision are less acute,
suggestive of its recent forest origins.
Cape buffalo are both diurnal and nocturnal; little is known about
65-85% of a 24-hour period is spent grazing and ruminating.
Grazing activity may total 5.3-13.4 hours/day, about equally spent
between night and day, though some populations may feed more at night.
Grazing peaks occur between 0600 and 1000 and from 1400 to 1800 h;
nocturnal grazing mostly occurs from 2000 to 0330 h. They may restrict
grazing to dawn, dusk, and nighttime in disturbed areas.
Resting and ruminating peaks from 0100 to 0500 and from 1200 to 1600 h,
although shorter resting periods occur every 0.5-3 hours. Buffalo
prefer to rest in the open, except in extreme heat or when disturbed
Buffalo visit water at least once/day.
Average daily movement is 1.2-8 km. Breeding herds travel 2-3 times as
far per day as do bachelor herds, and "pathfinder"
individuals usually guide the herds. Buffalo toward the front of a
moving herd tend to be in better physical condition than those toward
Mud wallowing, possibly related to grooming, may also have a social
significance because in some populations it is done most frequently by
dominant bulls, generally during the heat of the day.
Gregarious and nonterritorial.
Herds are small stable groups or clans of presumably related cows
gathered in a nearly linear dominance hierarchy and adult and subadult
bulls that are ranked by age and physical condition (i.e., size and
Social life appears to be built around mother-calf relationship. Social
licking is believed to foster bonds between herd members and mother
and calf. An offspring accompanies its dam until about 2-3 years of
age, after which time, it often forms a subgroup with other young.
Upon maturity, females are believed to remain in their clans, although
they no longer closely follow their dam.
Males end their maternal bond at adolescence. Young bulls generally
keep their distance from adults, and at about 4 years of age, they
begin to leave the herd in bachelor groups during the dry season.
Males more than 10 years of age remain permanently separated from the
Herds vary in size according to season and locality (i.e., availability
of food and water); size and composition can change frequently. Herds
occasionally mingle, but the herd is a recognizable unit to its
Average herd sizes range from 3.7 to 590, but herds may number up to
ca. 3,000 (Savuti, Botswana). Larger herds usually occur in more open
areas (e.g., extensive floodplains and broad river valleys), and the
largest may form near the end of the wet season, when mating peaks.
Large herds usually contain males, females, and young; however, males
are often solitary or in bachelor herds of usually not more than a
dozen animals and rarely having as many as 50 individuals. Many adult
males leave breeding herds after the rutting season, a time that may
correspond with dry-season fragmentation of their habitat.
Forest buffalo live in smaller groups (up to eight individuals) than
Threat behavior, rather than aggressive fighting, is usually sufficient
to maintain linear hierarchy among adult bulls.
Body size, which increases with age, determines dominance among males,
who are dominant over all females. Males 8-10 years old are usually
strongest and do the mating; herd bulls keep their standing about 4-5
years. There is less aggression among males in old bachelor groups
than in breeding herds.
Agonistic behaviors include: high-horn presentation (threat), lateral
display, rubbing face or neck on ground, ground-horning with earth
tossing, horning bushes, rolling in mud/dust, wallowing in deep mud,
Fighting can involve charging with chin raised, ramming, and
front-pressing. Fights occur infrequently, generally between two
individuals close in dominance status, and are usually brief, though
violent and possibly fatal; heaviest animal wins. Bulls often
head-spar, a milder form of fighting that has the general appearance
of play, to establish dominance status.
Running in circles, prancing, and butting, which are commonly
associated with sparring.
Young play more frequently than adults.
Dominance/threat displays include agonistic behaviors listed above and
the following: low-horn presentation, stiff-legged walk, and head
tossing and hooking.
Defensive/submissive displays include: head-low/chin-out posture and
Many calls are similar to the lowing of domestic cattle but are usually
lower-pitched. Buffalo are also less vocal than cattle, and they
rarely call except when in a group.
Vocalizations include the following signals: to move, direction-giving,
water, position, warning, aggression, mother-to-calf, calf distress,
danger, and various grazing vocalizations.
Aggressive vocalizations consist of grunting and growling, and
submission is indicated by bellowing.
A calf bleats and the dam answers with a croak when the two lose
Role is unclear, but olfaction is likely important for detection of
predators and individual recognition.
Buffalo are often accompanied by cattle egrets, which feed on insects
flushed during grazing. Oxpeckers feed on the ectoparasites of
Their grazing and trampling of old grass opens up additional land for
more selective species.
Mortality during the period between birth and reproductive maturity is
55-70%. Factors affecting all age groups include intraspecific
competition for food and water, diseases, and predators (including
humans), although older animals are more susceptible to death from
starvation. Populations are mainly kept in check by carrying capacity
of their habitats. Some populations may be limited by competition for
food with elephant or wildebeest.
Lions and spotted hyenas are the main predators, but leopards and
crocodiles occasionally make kills. Elephants have reportedly killed
buffalo. Lions may be unsuccessful (even gored or trampled to death)
in their attempts to pull down buffalo. Buffalo attack lions, often in
mobs, when their calves are threatened. Herds react to predators by
bunching and confronting the predator with a solid mass of horns. Lone
bulls are more susceptible to predation, though are often in better
physical condition, than animals in breeding herds. In the absence of
epidemics or drought conditions, about 90% of buffalo die from
Native cultures regard buffalo horns as symbolizing virility or
fertility, and horns are commonly used in traditional witchcraft.
Buffalo are prized for their meat in many countries.
Domestication attempts have largely failed due to their intractable
Buffalo are classified as bulk and roughage eaters that are dependent
Grasses and sedges form the bulk of the diet, but the leaves and shoots
of other plants (e.g., shrubs, trees, and herbs) are also eaten. Diet
may be determined solely by availability; however, when a choice
exists, plants high in protein and carbohydrates may be selected to
maximize nutrient intake.
The major elements aluminum and iron may be the main chemical stimuli
for soil-eating behavior.
Less prone to selective overgrazing than wildebeest and zebra and less
destructive and more economical than most other grazers in utilizing
available food, including old grass.
During their nomadic foraging routines, they range up to 18 km from
their watering places, which they visit at least once (and often
twice) a day. Buffalo cannot tolerate water restriction for very long
during hot temperatures. Adults need up to 39-55 liters/day. Water
consumption may be 30-40 liters/day; the rest is obtained via food and
As is typical of grass-eating species, the ruminant stomach is
highly-differentiated and subdivided in order to utilize bulky food
rich in cellulose. Rumen compartments hold ingested food for microbial
Daily food intake (dry forage) averages 6.1-17.5 kg.
Estrus cycle averages 23 days. Female is in estrus 1-6 days during this
period. Although there may be several consecutive cycles during the
breeding season, it is uncertain if females cycle throughout the year.
Bulls routinely check estrus status of cows by flehmen (urine-testing),
and they can stimulate females to urinate by licking the vulva.
A bull will try to keep other bulls away from a cow in estrus, but his
possessiveness and her evasiveness usually attracts other bulls,
leading to a sequence of displacements by more dominant bulls. An
estrus female may mate with this succession of bulls, but dominant
males obtain most of the matings.
Female's receptiveness to mating is tested by the attending bull as he
licks and rests his chin on her rump; if she stands, he mounts. A cow
will stand still with tail arched when she is ready for copulation.
cows do not calve every year, and lactational anestrus apparently
follows parturition; there is little sexual activity during this time.
Interbirth intervals are 15-36 months; intervals tend to decrease as
Birth rate probably varies in relation to food availability.
Seasonal births coincide with the period of optimum grass growth; less
stress is experienced if lactation occurs during the rains.
Non-seasonal breeding occurs in areas where environmental conditions
are favorable year-round (Happold,
1987). In captivity, buffalo breed
throughout the year (Gingerich,
Calving occurs when the herd is resting (i.e., during the afternoon or
before dawn). The dam remains with or near the herd, but she and calf
may be briefly left behind until the calf gains strength. The newborn
is underdeveloped compared with calves of other grazing ungulates.
Although on its feet within 10 minutes of birth, it is too weak to
follow for several hours and is a slow runner for several weeks.
Neonates are occasionally hidden in brush when the dam moves away to
Body weight is 32-50 kg (females) to 26-54 kg (males); mean is
estimated to be 40-45 kg.
Deciduous teeth usually consist of six incisors on the lower jaw and
four to six premolars on both the upper and lower jaws.
Body length is 121-150 cm (females) to 132-151 cm (males).
Horns are straight and appear V-shaped when viewed frontally.
Individual coloration is variable, as it is in adults, but a dark
neonate often lightens a few months after birth; thereafter, it tends
to darken with age.
Calves are weaned at 4-6 months, but in some populations, suckling may
last 18 months or until the next calf is born. They graze for brief
periods at about 2 months of age.
Juveniles (1-2 years of age):
Body weight is 259-323 kg (females) to 283 kg (males).
Body length is 178-196 cm (females) to 194 cm (males).
Horns are 30-46 cm long and curve slightly outwards.
Deciduous dentition is complete: 0/3, incisors; 0/1, canines; 3/3,
premolars; total, 20. It may be complete as early as 3 months of age.
Subadults (2-3 years of age):
Body weight is 314-493 kg (females) to 282-483 kg (males).
Body length is 191-219 cm (females) to 191-225 cm (males).
Horns are 41-86 cm along the curvature, and the tips grow towards one
another. In bulls, horns thicken at their bases, and the boss begins
to take shape.
Tooth replacement is at a maximum in the 3-5 year age groups.
Males can reach sexual maturity at 2.5-3 years of age, but older herd
bulls usually prevent them from breeding until they are at least 7-8
years of age. Age of puberty is determined by body weight.
Young adults (4-5 years of
Body weight is 397-576 kg (females) to 378-617 kg (males). Females
attain adult weight during fourth year, when they are also sexually
mature and usually give birth to their first calf.
Males maintain growth throughout their lives.
Body length is 215-238 cm (females) to 211-255 (males).
Tips of the horns begin to sweep backwards. Horns reach their adult
shape in cows; the boss (i.e., the raised, rounded area at the base)
continues to enlarge in bulls.
Adults (>5 years of age):
Body weight is 312-637 kg (females) to 488-768 kg (males).
Body length is 215-249 cm (females) to 225-267 cm (males).
In older cows, tips of the horns are widely separated, and they wear
and may break with age. In bulls, the backward sweep of the horns is
completed, and the boss is well-shaped, though it continues to thicken
with age. The widest span is reached in this age class (up to 104 cm),
but the length of the horns along the curvature can be less than in
younger age classes.
Coloration may change to black or dark brown; old animals become nearly
hairless in spots, and hairs may become white, particularly those on
the face and neck.
Permanent dentition is in place by 6 years of age.
Longevity:20-25 years in nature; 29 years,
6 months in captivity (Jones,
IUCN status for savanna buffalo is Lower risk/conservation dependent,
and the forest buffalo is listed as Lower risk/near threatened; not
listed by CITES.
Although the African buffalo is not in danger of extinction,
populations in the mountains of western and central Africa are rapidly
declining. The current wild population of savanna buffalo is estimated
to number approximately 500,000-1,000,000; forest buffalo number about
60,000. The population was estimated to be 3-4 million during the
early 1950s (Blancou,
1958) and 2-3 million in the early 1970s (Best
and Raw, 1975).
Buffalo can increase their numbers quickly in protected areas, and may
need culling to avoid habitat degradation caused by overpopulation (Estes,
1991). The population in the 2050 sq km Yankari Game Reserve,
Nigeria, increased from an estimated 296 individuals in 1963 to 680 in
Recent population estimates in protected areas are as follows: Angola,
<500; Benin, >2,000; Botswana, 8,050 (plus 18,840 in other
areas); Burkina Faso, 1,620; Burundi, 500; Cameroon, 3,210; Central
African Republic, 19,000; Chad, 1,020; Congo (Zaire), 39,180;
Ethiopia, 2,330; Ivory Coast, >8,330; Kenya, > 11,630 (plus
7,930 in other areas); Malawi, >3,150; Mali, 120; Mozambique,
9,570; Namibia, 690 (plus 310 in other areas); Niger, 500; Nigeria,
>200; Rwanda, 1,200; Senegal, 4,000; South Africa, 28,470 (plus
>2,500 on private land); Sudan, >100; Tanzania, >245,000
(plus 97,350 in other areas); Uganda, >20,220; Zambia, >40,090
(population for entire country); Zimbabwe, >50,330 (population for
Population densities of savanna buffalo in African reserves were:
Ngorongoro (1964-1968), 0.2/sq km; Kruger (1954-1966), 0.6/sq km;
Serengeti (1971-1972), 6.8/sq km; Albert (1956), 13.9/sq/km; and
Manyara (1967-1969), 16.5/sq km (Delany & Happold,
1977). Average density of savanna buffalo was
estimated to be 0.05-0.6/sq km. Forest buffalo densities were
0.01-7.4/sq km, and average density was estimated to be 0.03-0.3/sq
Threats to survival:
Habitat loss resulting from agricultural development and competition
from domestic cattle.
Diseases, particularly those transmitted from domestic livestock.
First zoological collection known to exhibit African buffalo was London
St. Louis Zoo was the first in North America to exhibit the species
San Diego Zoo received 1.1 Cape buffalo from St. Louis in 1946; the
first calf born in 1949. 1.0 forest buffalo received from Bronx Zoo in
1968, and the first birth occurred in 1980 at the Wild Animal Park.
Management and husbandry considerations are addressed by Kleiman et al.
(1996) and Shoemaker