African buffalo

Subspecies

Much discussion has been made about the African buffalo's taxonomy because throughout its range, populations differ noticeably in physical appearance. The IUCN Red List currently lists four subspecies: the Cape buffalo (S.c. caffer), the African forest buffalo (S.c. nanus), the West African savanna buffalo (S.c. brachyceros), and the Central African savanna buffalo (S.c. aequinoctialis). Some authors have proposed that only three subspecies—the Cape, African forest, and northern savanna buffaloes (S.c. umarii)—would be recognized if genetic evidence is taken into account when determining subspecies, while others have suggested that the forest, and West and Central African buffaloes be classified as a single subspecies because of their phylogenetic similarities. However, studies of the genes of populations in different regions provide no support to either conclusion.

Results derived from maternally inherited mitochondrial DNA and paternally inherited Y-chromosomes reveal two lineages of African buffalo. The first lineage is made up of a single subspecies, the Cape buffalo, which is found in East and Southern Africa, while the second includes West and Central African populations, of which there are three subspecies. Spatial patterns of microsatellites show that the Cape buffalo lineage, as well as the West and Central African subspecies, underwent slow, continuous genetic changes throughout their ranges. Based on these patterns, there was once a large metapopulation of savanna buffalo. In the Central African Republic, the forest buffalo shows similar clinal patterns, but with double the genetic changes accumulated possibly due to restricted gene flow and the fact that there are less forest buffaloes per square kilometer than savanna buffaloes.

The following is a table based on the taxonomic classification used by the IUCN.

Name	Image	Description	Ecological grouping
Cape buffalo (S. c. caffer)		The largest and most abundant subspecies, with males weighing up to 900 kg (2,000 lb). Horns can grow to a maximum of 165 cm (65 in) in males and 150 cm (59 in) in females. Adults are usually black in color and have a skin thickness of more than 2 to 3 cm (0.79 to 1.18 in). In males, the large skull is held up by a hump situated above the shoulders.	Savanna buffalo
West African savanna buffalo (S. c. brachyceros)		Adults are 120 to 145 cm (47 to 57 in) tall at the shoulders, 200 to 245 cm (79 to 96 in) in body length and weigh 300 to 600 kg (660 to 1,320 lb). Reddish individuals are found across the subspecies' range. Horns have a length of 56 to 103 cm (22 to 41 in) and a span of 56 to 103 cm (22 to 41 in). In some parts of its range, there have been reports of hybrids with the forest buffalo.	Savanna buffalo
Central African buffalo (S. c. aequinoctialis)		Intermediate subspecies similar in physical dimensions to the West African buffalo. Red forms exist across its range.	Savanna buffalo
African forest buffalo (S. c. nanus)		The smallest subspecies, with a body mass about 50% smaller than that of the Cape buffalo. The coat is rusty to black in color, and horns are up to 69 cm (27 in) long. Compared to other subspecies, adults have less pronounced sexual dimorphism.	Forest buffalo

Social behavior

Adult females form the core of African buffalo herds. The number of individuals in herds is variable and differs across regions; for West and Central African buffaloes, herds of females and their young have an average of 45 and a maximum of 150 individuals, whereas in Southern and East African floodplains, Cape buffaloes are known to form large herds of up to 2,000 members. About 3–25 animals make up a forest buffalo herd.

Female savanna buffaloes remain with the herd for life and may continue bonding with their mother even after the age of weaning. On the other hand, young males grow increasingly detached from their mothers and separate from the herd to join bachelor groups at age four to five for a limited period. They later join a herd to compete for mating opportunities. As males lose their strength, they again leave for bachelor groups so as to recover by excessively grazing in areas inaccessible to herds. The permanent departure of males aged ten or older from mixed herds has been recorded and appears to happen randomly. In the forest buffalo, males make up the minority of the stable herd and travel with the group throughout the year.

Savanna buffaloes form durable groups that have been known to last over 50 years. These groups generally show a strong preference for their home ranges. In mixed herds, fission-fusion behaviors exist, where members briefly separate and form subgroups, then unify once more. Separation can occur during both the wet and dry seasons. Sometimes, separation may be due to large herd size. A few bulls and some females lead the herd when foraging. Decisions on where to eat or drink are not made by these individuals but rather through a process called 'voting.' After a lengthy resting session, right before dusk, adult females will stand up, raise their heads, and point in a particular direction, then again rest. If multiple voting females point in the same direction, consensus is strong. However, in cases of weak consensus, the herd may separate and rejoin days later.

Most forest buffalo sightings are of groups, though solitary individuals of both sexes have been observed. Solitary male sightings are more frequent than sightings of solitary females. In Dzanga-Ndoki National Park between 2007 and 2008, short-lived fission-fusion dynamics and a correlation between an increase in herd size and fission-fusion dynamics were recorded in a specific herd. Another finding was that during the wet season members of the herd traveled farther away from each other in smaller groups.

In savanna buffaloes, sparring occurs in both subadult and adult males, often concluding with hooking of the other buffalo by the victor after a period of horn butting and shoving. Possible reasons for sparring include the assessment of dominance or enjoyment. Subadult females rarely interact with each other and when they do, interactions are often limited. Positioning within a herd is observed in adult females and their young. Females may inherit a specific location from their mother. Behaviors such as using horns for butting and hooking of opponents on their side and hindquarter may contribute to positioning within a herd. In some areas, male buffaloes have a pecking order. Dominant individuals accentuate their shoulders during a threat posture by lowering the nose while lifting the head up. Often while bellowing, subordinate males respond to challenges by lowering the head and moving behind a dominant male to position the nose between his hindlegs. Fighting among males is extremely uncommon as in most cases, males observe each other from a distance and avoid direct interaction. Only when two bulls are evenly matched do fights occur, and such encounters could turn deadly. Although submission has been recorded in subadult males, adult females and juveniles in Ika Bai, Cameroon, forest buffaloes display few aggressive or dominance-related behaviors when observed. During feeding and resting, most interactions between herd members according to a 2007 study in Dzanga-Ndoki National Park, involves the chasing, usually by females, of other other group members, usually juveniles. Intraspecific competition for ideal spots is cited as a possible cause.

The African buffalo rarely makes sounds such as grunts, beeps, bellows, snorts, croaks and coughs. When seriously or mortality injured, a bull may emit a loud bellow which could attract the attention of another bull who assists in about 33% of cases involving predator attacks. Coughs and snorts are usually made when threatened. Conversely, buffalo remain silent, particularly at night, in order to more easily spot threats and warn others, though no alarm call exists.

Habitat use and activity patterns

Home ranges for herds of savanna buffaloes vary from as small as 50 km² (19 sq mi) to as large as 350 km² (140 sq mi). Forest buffaloes have the smallest home ranges of any subspecies, and are mostly found in forests during the dry season and marshes during the wet season. In savanna populations, herds that have to travel large distances seasonally to locate patchily distributed resources have greater home ranges. Bachelor groups on the other hand, have small home ranges of only 0.5 to 4.0 km² (0.19 to 1.54 sq mi). Herds in some environments increase their home ranges during the wet season as resources become plentiful, but are restricted to the vicinity of water sources during the dry season. In water-rich environments, home ranges may instead expand during the dry season.

Migration has been observed in savanna populations. Throughout their range, they do not always follow the same migration patterns and in some areas, not every member of a group may choose to migrate. According to a 2014 study, the longest African buffalo movements occur in the Kavango–Zambezi Transfrontier Conservation Area, including one case of a female which undertook a more than 170 km (110 mi) round trip in successive wet seasons. Most migrations happen during this period because of the availability of good quality grass in savannas and upland woodlands. The curing of grass and the retreat of water in these habitats during the dry season eventually forces buffaloes to migrate to lowland areas as they are home to moist soil for vegetation growth. West African buffaloes in W Regional Park have been recorded in a 2011 study undertaking longer migrations from a permanent water source during the wet season because vegetation became more plentiful in south-west areas. Herds moved 25 to 45 km (16 to 28 mi) south-west from permanent water sources during the monsoon, whereas in the dry season, they remained within 3.3 to 5.3 km (2.1 to 3.3 mi) of one.

The African buffalo spends about 17 to 19 hours a day feeding and ruminating. It is highly dependent on water, requiring 45 L (9.9 imp gal; 12 U.S. gal) daily every 2 or more days. Feeding generally occurs from dawn to mid-morning hours and from late-evening to mid-night hours. The total feeding time during the night is comparable to the feeding time during the day, and across seasons, feeding time is greater in the dry than in the wet season. A study conducted in West Africa in 2011 found buffaloes to spend equal amounts of day and night hours performing daily activities, especially during crepuscular hours.

Savanna buffalo populations have been recorded averaging less than 4 km (2.5 mi) or more than 6 km (3.7 mi) of distance traveled per day. On the contrary, forest buffaloes only travel 0.5 to 1.5 km (0.31 to 0.93 mi) per day, according to a study in Dzanga-Ndoki National Park, Central African Republic. During the late afternoon, when temperatures are at their highest, forest and male savanna buffaloes wallow to keep cool, though the former wallow more during the dry season. Members of mixed herds in savanna populations typically seek shade from the searing midday heat.

Diet

As a ruminant, the African buffalo derives most of its diet from grass and dietary fiber; per day, it consumes 6.5 to 15 kg (14 to 33 lb) of vegetation. In some cases, it is known to live off grass that most other herbivores either cannot reach or chew, though it is unable to shorten grass as much as other herbivores and instead clears the landscape for short-grass herbivores, thereby filling the ecological niche of an ecosystem engineer. Its preferred foraging grounds include floodplains and forest glades, as they offer large pastures and grass leaves of at least 10 cm (3.9 in) tall. The plains zebra (Equus quagga), African bush elephant (Loxodonta africana, blue wildebeest (Connochaetes taurinus), and cattle (Bos taurus) compete with the African buffalo for resources.

The Cape buffalo subspecies prefers grass such as panicgrass, Guinea grass, and fonio, and during the dry season when food loses much of its nutrition, vital energy sources include Leersia hexandra, Setaria sphacelata, and Cyperus laevigatus. In West and Central African buffaloes, perennial grass, beard grass, foxtail, and flatsedges are the preferred grass genera. During the dry season, as grass turns woody in texture, savanna populations partially adopt browsing herbivory, consuming shrubs and tree leaves. Tall grasses and sedges, as well as reeds, are foods that savanna populations subsist on during droughts because they are easier to digest and are more nutritious. Studies on forest buffalo feeding ecology suggest that it feeds on graminoids, arrowroot, day flowers, and cryptogams.

Reproduction

Most Cape buffalo calves are born during the wet season, except in more mesic areas. Data on forest buffalo reproduction is limited, though observations indicate that births can occur during both the dry and wet seasons. Female Cape buffaloes in captivity enter heat for 5 to 6 days in a 23-day cycle, according to one study. Males assess a female's readiness to mate by sniffing chemicals in her urine. Usually, dominant males mate with most of the females in heat. After a gestation of around 340 days, females typically give birth within the herd to a single calf, weighing 30 to 40 kg (66 to 88 lb). Calves nurse from the hindquarters, which allows the mother to move with the herd while continuing to suckle her young. As calves increase in age from one to six month(s), they decrease the time spent suckling by 50%; weaning occurs between 10 and 18 months, depending on factors such as population density and competition between buffaloes. Females begin breeding by age five, which is less than two years after they start to ovulate, while males typically breed by at least age seven, despite being sexually mature at five to six. Lifespan is about 12 years in the wild, though some individuals may reach 20. In captivity, the maximum lifespan is nearly 30 years.

Female forest buffaloes give birth to a single calf every two years after a gestation of more than 300 days. A 2008 study in Dzanga-Ndoki National Park observed that during birth, females leave the herd for a few days and when feeding, hide their calves in vegetation. Similar behaviors have been observed in Cape buffaloes. Once calves are able to walk steadily, female forest buffaloes return to the herd and raise their young for two or more years. Both sexes reach adulthood past the age of five and females first breed by at least age three.

Mortality factors

• 
  Decomposing skull
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  Male lion and cub feeding on a Cape buffalo carcass

When triggered, the African buffalo becomes aggressive, a behavior that along with its large size ensure that it is a target for only a few predators; calves, on the other hand, are vulnerable to predation by multiple predators, including lions (Panthera leo), African wild dogs (Lycaon pictus), spotted hyenas (Crocuta crocuta), leopards (Panthera pardus), and Central African rock pythons (Python sebae). Nile crocodiles (Crocodylus niloticus) are known to prey on adult savanna buffaloes, though lions are the main predators of the Cape buffalo subspecies. To sneak up on a herd and isolate an individual, lions typically take advantage of thick vegetation. If successful, lions attack the rear to bring a buffalo down but because of the buffalo's thick neck the felines are instead forced to subdue the animal with a suffocation bite to the nose. In other cases, the buffalo drives off or even kills lions. By positioning its rear to thick bushes and utilizing its horns, a buffalo becomes especially successful at fending off lions and in open areas, large herds may offer protection from lion predation. Buffalo can also leap around 1.5 m (4 ft 11 in) and run at top speeds of 55 to 60 km/h (34 to 37 mph). In exceptional cases, the forest buffalo has been known to fall prey to leopards. Possible anti-predator behaviors of this subspecies include resting and moving as a group, and stampeding characterized by a female taking the lead and a male trailing behind.

Like all ungulates, the African buffalo is likely susceptible to rinderpest, a disease caused by Morbillivirus. Transmission occurs when respiratory droplets or feces from an affected animal contaminate a healthy one. Severe diarrhea and dehydration are associated symptoms that often lead to death. The African buffalo serves as a maintenance host for Mycobacterium bovis, a bacteria that causes bovine tuberculosis. Other infections include brucellosis, anthrax, haemorrhagic septicaemia, foot-and-mouth disease, theileriosis, trypanosomiasis, rabies, tick-borne diseases, and African insect-borne diseases.

Hunting

Prior to colonialism, the African buffalo served as an important source of protein for African societies like the Valley Bisa of Zambia, who regarded buffalo meat as integral to their identity and fate. In Ethiopia, the African buffalo is poached to increase one's social ranking, and among western hunters it is considered a major animal for trophy hunting due to the danger it poses during such activities. Out of 37 range countries, the African buffalo can be legally hunted in 16 of them as of 2022. Buffalo-caused human injuries or death have been reported and seem to happen regularly.

In the late 19th century, a rinderpest outbreak killed 95% of African buffaloes, and targeted hunting by foreign settlers led to further population declines. By the early 20th century, the African buffalo received protection from uncontrolled hunting. During the same period, safaris and hunting for sport became more popular throughout Africa. Starting in East Africa and expanding after World War I, hunting safaris became dictated by law and after World War II they became increasingly structured as a business. A rise of hunting safaris in other African regions occured following the 1970s permanent and temporary hunting bans in Kenya and Tanzania, respectively.

In human culture

Humans and African buffaloes have co-existed for thousands of years. Across African communities and around the world, the African buffalo contributes to human culture. The earliest cultural depictions of African buffalo in Africa are dated to the Middle Stone Age androck paintings depicting this animal have been observed in San culture. Although it is positively affiliated in Ghana with the word "uprightness", it is described by hunters and African people as being wild, dangerous, powerful, intimidating and vengeful. In some African societies, the species is used as a totemic symbol of strength, courage, danger and potent spirituality. Masks of buffalo and similar species, which may signify a "bush cow", have been observed in West African tribes; crosses between animal, human and spirit are sometimes represented with these masks which are used mainly for dancing and rituals during male initiation ceremonies. The progression from the natural world to human settlements is particularly signified by formidable cow masks and it appears that many rural societies in Africa have folklore about where livestock came from that place cattle close to humans and buffalo to nature. In North Africa, the Berber have a legend stating that the ancestry of all wild animals and cattle can be traced back to an African buffalo and a heifer, respectively. Analogously, in South Sudan, Nuer groups credit the African buffalo for being the biggest killer of people, and describe the species as being vengeful towards people because it was people who slayed the common mother it shared with the cattle. It is also believed that cattle avenge this loss by first being a part of the human world then by prompting people to fight and kill each other over them.

Many African cultures forbid people from invading a buffalo's space or using its body parts, including ethnic groups of southern Africa and Cameroonian pygmies, who do so for totemism and protection against demons, respectively. On the contrary, shamans and traditional healers commonly use buffalo body parts to cure ailments and disease, improve one's sexual life, and fight off demons.

The African buffalo has been used as an icon in numerous social domains, including sports, military units, merchandise, and shipping. The Buffalo bicycle is an African buffalo icon that is 'built for big loads on tough roads in Africa,' a slogan highlighting its size and strength.

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