African buffalo

Extant subspecies

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 head and horns are carried 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 weight range between 265 and 320 kg (584 and 705 lb), about half the size 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

Evolution

Syncerus and Bubalus (Asian buffaloes) split from a common ancestor around 6—9.8 million years ago, according to genetic evidence. S. acoelotus was the earliest known species of Syncerus and was equal in size to the Cape buffalo but with smaller horns. Currently, the African buffalo is the only extant species of the genus, emerging 500 thousand to 1 million years ago. It spread across sub-Saharan Africa during glacial and interpluvial periods as conditions allowed for savanna expansion. Around 130–300 thousand years ago, drastic environmental changes isolated populations into savanna refugia, leading to the formation of two genetic clusters. During the late to middle Pleistocene, the species again dispersed from Central Africa, with one cluster spreading across West Africa, while the other reached East and Southern Africa. Genetic evidence indicates that the forest buffalo arose before the Cape buffalo.

Social behavior

African buffalo herds have a core comprising adult females and their young. The number of individuals in a herd is highly variable and depends on the subspecies and region. In Southern and East African floodplains, the Cape buffalo is known to form the largest herds of any subspecies, as numbers tend to exceed a thousand. In contrast, only 3–25 animals make up the average forest buffalo herd. For West and Central African buffaloes, typical herd size ranges from 45 to 150 individuals.

Female African buffaloes remain with the herd for life and, in some subspecies, maintain a close bond with their mother post-weaning. On the other hand, aside from the forest buffalo, young males become less and less dependent on their mothers and separate from the herd to temporarily join bachelor groups at age four to five. After a few years, they 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. In the forest buffalo, males make up the minority of the stable herd and travel with the group throughout the year.

Mixed herds are fission-fusion societies, where members briefly separate and form subgroups, then unify once more. Separation seems to occur more during the wet season, when there is a resource surplus. 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 will separate and rejoin days later.

There is a pecking order among male buffaloes. Dominant individuals accentuate their frame during a threat posture by lifting the head up and pointing the nose down. Subordinate males respond by lowering the head and positioning the nose between a dominant bull's hindlegs; they often bellow throughout the interaction. In most cases, bulls 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.

The African buffalo is mostly silent but can make sounds such as grunts, bellows, snorts, and coughs. When injured by predators, a bull may emit a loud bellow to signal for help. Coughs and snorts are usually made when running or when a predator is spotted, though no alarm call exists.

Habitat use and activity patterns

African buffalo herds have home ranges that vary from as small as 4.6 km² (1.8 sq mi) to as large as 350 km² (140 sq mi). Forest buffaloes have the smallest home ranges of any subspecies, and habitat use depends on the season, as herds prioritize forests in the dry season and marshes in the wet season. In savanna populations, herds occurring in areas where resources are widely distributed have the largest home ranges and often migrate seasonally. 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). Field observations reveal that herds in some environments increase their home ranges in the wet season as resources become plentiful and decrease them in the dry season because of a water shortage that forces herds to settle in areas near a viable water source. In contrast, during the dry season in environments with plenty of viable water, herds will expand their home ranges as they search for ideal feeding grounds.

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) every 48 hours. 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. In most populations, activity appears to occur during both day and night. 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 average 4 to 6 km (2.5 to 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 mostly male savanna buffaloes wallow to keep cool, though the former wallow more in the dry season. For thermoregulation purposes, members of mixed herds in savanna populations typically seek shade from the searing midday heat.

Diet

The African buffalo is a ruminant that primarily feeds on grass and dietary fiber. Per day, it consumes 6.5 to 15 kg (14 to 33 lb) of grass, about 2.2% of its body weight. In some cases, it is known to live off grass that most other grazers either cannot reach or chew. However, it is unable to shorten grass as much as other herbivores and instead clears the landscape for short-grass grazers, thus 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 Cape buffalo subspecies prefers grass such as panicgrass, Guinea grass, and fonio, and during the dry season when food is scarce and of poor nutritional quality, species such as Leersia hexandra, Setaria sphacelata, and Cyperus laevigatus become important food sources. In West and Central African buffaloes, perennial grass, beard grass, foxtail, and flatsedges are the preferred grass genera. During the dry season, as grass lignifies, the savanna population partially adopts browsing herbivory and also consumes shrubs and tree leaves. Studies on forest buffalo feeding ecology suggest that it mainly 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. A female enters heat for 5 to 6 days in an 18- to 23-day cycle. 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 340 days, females typically give birth within the herd to a single calf, weighing 30 kg (66 lb). Calves nurse from the hindquarters, which allows the mother to move with the herd while continuing to suckle her young. As calves get older, they spend less time suckling, from around 10 minutes at one month to 5 minutes by six months; weaning occurs between 10 and 18 months. Females begin breeding by age five, which is 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.

Mortality factors

Adult African buffaloes have few natural predators, owing to their aggressive temperament and large size. Calves, however, are much more vulnerable to predation and require constant protection. The lion (Panthera leo) is the major predator of the Cape buffalo. A pride typically takes advantage of thick vegetation to sneak up on a herd or bachelor group and isolate an individual. If successful, pride members attack the rear to bring the buffalo down and then subdue it with a suffocation bite to the nose as opposed to the neck, which is too thick to break. In other cases, the Cape buffalo drives off or even kills lions. It is especially successful at fending off lions in thick bushes, where it faces them head-on while positioning its rear in the thickets. In open areas, large herds may offer protection from lion predation. Spotted hyenas (Crocuta crocuta) and leopards (Panthera pardus) are minor predators of the Cape buffalo that mostly target calves. On rare occasions, the forest buffalo has been known to fall prey to leopards. Possible anti-predator behaviors of this subspecies include communal resting and movement, as well as stampeding characterized by a female taking the lead and a male lagging the last behind.

Like all ungulates, the African buffalo is susceptible to rinderpest, a disease caused by Morbillivirus. Transmission occurs when respiratory droplets or feces from an affected animal come into close contact with 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 and Zambia, the African buffalo is sometimes poached to increase one's social ranking, and among western hunters it is considered a major animal for trophy hunting due to its aggression and power. Out of 37 range countries, the African buffalo can be legally hunted in 16 of them as of 2022. Attacks on humans have been reported and can cause serious injuries or death.

In the late 19th century, a rinderpest outbreak killed 95% of African buffaloes, and targeted hunting by European 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 II, hunting safaris became increasingly structured as a business and were dictated by law, and following the 1970s permanent and temporary hunting bans in Kenya and Tanzania, respectively, other African regions experienced a rise in hunting safaris.

In human culture

Humans and African buffaloes have co-existed for thousands of years. Across African communities, the African buffalo contributes to human culture, and rock paintings depicting the African buffalo have been dated to as far back as the Middle Stone Age. The local African people generally describe the species as being a dangerous and powerful animal that defends itself viciously. In southern and West Africa, the African buffalo is widely used as a totemic symbol of strength, courage, and potent spirituality. Horizontal masks depicting the African buffalo have been observed in West African tribes. Many of these masks represent a cross between animal, human, and spirit and are mainly used for dancing and rituals during male initiation ceremonies. Further north, 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. In South Sudan, Nuer groups credit the African buffalo for being their biggest cause of mortality, and describe the species as being vengeful towards humans because a group of people slayed the common mother it shared with the cattle; it is also believed that cattle avenge this loss by prompting people to fight and kill 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 routinely 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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