The African buffalo (Syncerus caffer) is a large bovid native to sub-Saharan Africa. First described in 1779 by Swedish naturalist Anders Sparrman, it is the only extant species of the genus Syncerus. Four subspecies are currently recognized: the Cape buffalo, West African savanna buffalo, Central African savanna buffalo, and African forest buffalo. It is a heavily built animal with a broad head and torso, a strong neck, large horns, and a bony plate or "boss" between the horns of males. The fur ranges in color from rusty to black, and the tail ends with a dark, hairy tuft.
The species occurs in a variety of habitats, such as woodlands, savannas, forests, grasslands, shrublands, and wetlands. It is a ruminant and a grazer, primarily feeding on grass and dietary fiber. A social animal, the African buffalo forms herds ranging in size from a few individuals to over a thousand members. Herds typically consist of adult females and their young, and a number of adult males that temporarily join herds for mating opportunities. Males have a pecking order where dominant individuals hold the best access to mates. After a gestation of 340 days, females give birth to a single calf. Weaning occurs between 10 and 18 months, and the lifespan is about 12 years in the wild. Adult African buffaloes have few natural predators because of their size and strength, though calves are much more vulnerable.
Since the 19th century, the African buffalo's geographic distribution has greatly reduced due to human factors such as poaching, climate change, agriculture, and disease outbreaks. Despite this, conservation efforts have managed to increase populations in some parts of its range. As of 2019, it is listed as Near Threatened on the IUCN Red List, with an estimated population of 398,000 to 401,000 mature individuals.
Taxonomy
The African buffalo was first described by Anders Sparrman in 1779. Due to its pronounced polymorphism, there has been much discussion about its taxonomic status. The IUCN Red List currently lists four subspecies: the Cape buffalo (S.c. caffer), the African forest buffalo (S.c. nanus), the West African buffalo (S.c. brachyceros), and the Central African buffalo (S.c. aequinoctialis). A 2013 study on maternally inherited mitochondrial DNA revealed two genetic clusters of African buffalo. The first cluster 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. These results were corroborated by a study on both paternally inherited Y-chromosomes and maternally inherited mitochondrial DNA.
Spatial patterns based on microsatellites show that the Cape buffalo cluster, as well as the West and Central African subspecies, underwent slow, continuous genetic changes in relation to an environmental gradient. Based on these patterns, it is believed that savanna buffalo populations (the Cape, West, and Central African buffaloes) were once part of a large metapopulation. In the Central African Republic, the forest buffalo shows similar clinal patterns, but with double the genetic changes accumulated. Low population density and restricted gene flow are likely causes for these differences.
It has been proposed that if genetic evidence is used as an extra criterion for determining subspecies, only three would be recognized: the Cape buffalo (S.c. caffer), African forest buffalo (S.c. nanus), and northern savanna buffalo (S.c. umarii). In 2014, a study suggested that the forest, and West and Central African buffaloes be classified as a single subspecies due to their phylogenetic similarities.
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.
Habitat
Historically, the African buffalo ranged throughout most of sub-Saharan Africa in habitats with yearly rainfalls above 250 mm. Since the 19th century, geographic distribution has greatly reduced due to human factors such as poaching, climate change, agriculture, and disease outbreaks. Most savanna buffalo populations are restricted to protected areas and nearby hunting zones. The Cape buffalo occurs across East and Southern Africa from Ethiopia southwards to South Africa and westwards to Namibia. Both the West and Central African buffaloes are found within the Sudano-Sahelian zone, with the former ranging from Senegal in the west to the Central African Republic in the east, whereas the latter is distributed north to Chad, south to the Democratic Republic of Congo, and east to Ethiopia. Two separate populations of forest buffalo reside in West and Central Africa, respectively.
The African buffalo can be found in a variety of habitats, such as woodlands, savannas, forests, grasslands, shrublands, and wetlands. It prefers semi-arid environments with a permanent water source not beyond . In East and Southern Africa, buffalo populations prefer woodlands, while those in West and Central Africa are found in shrub savannas and Sudanian woodlands, among other habitats. The forest buffalo is primarily found in forest-savanna mosaics in elevations of up to .
Description
The African buffalo is a large bovid characterized by its stocky build with a broad head and torso, thick, muscular neck, stubby legs, and curved hooves that grow larger in the forelegs than the hindlegs. The sagged, often scarred ears are outlined with long hairs, and the broad muzzle has a wet nose and wide eyes. The African buffalo is rusty to black in color and has a dark, hairy tuft at the end of the tail. Before obtaining the adult coat, young Cape buffaloes change in color from tawny to dark brown, and gradually lose their coat thickness. Adult forest buffaloes typically have a rusty to black coat pattern, whereas the coat of young individuals is denser and more reddish, sometimes with black markings.
Both sexes have large horns; however, males possess a fusion of bone called a boss between the bases of their horns. The boss is most pronounced in male Cape buffaloes. Females of this subspecies have smaller horns with hairy skin between them, and like their male counterparts, their horns curve laterally. Compared to the Cape buffalo, West and Central African buffaloes possess similar laterally curved horns, but without the downward arch. Horn span measures between and in savanna buffaloes. For forest buffaloes, horn morphology is described as being pointed horizontally backwards, lacking a boss, and reaching a length of up to .
The African buffalo is among the most polymorphic of African mammals, as subspecies vary considerably in size, color, and horn shape. The largest subspecies, the Cape buffalo, has a head-to-body length of and a shoulder height of . Males weigh , whereas females reach a maximum of . At tall at the shoulder, in body length and in weight, the forest buffalo is the smallest subspecies. The tail length, though differing across subspecies, is .
Behavior and ecology Social behavior
African buffalo herds are largely matriarchal, with the 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 eastern 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.
Observations on savanna buffalo social dynamics show that both mixed herds and bachelor groups exhibit stable membership and the preference for a certain home range over long periods of time. 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 to as large as . 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. Conversely, bachelor groups have small home ranges of only . The home ranges of mixed herds in some ecosystems has been observed increasing in the wet season because of the influx of resources across the land and decreasing in the dry season because the lack of water forces herds to settle in areas near a viable water source. Furthermore, during the dry season in ecosystems with plenty of water, herds will expand their home ranges as they search for ideal feeding grounds.