Syncerus cafferAfrican buffalo

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Geographic Range

Historically, African buffalos were found throughout most of sub-Sahara Africa. Today their distribution is considerably reduced and in many areas largely limited to reserves. Their habitat is fragmented in many areas by human activity and their numbers have been hugely reduced since the late 1800s from the introduced rinderpest epidemics, resulting in extirpations. African buffaloes are still distributed throughout Africa, and different portions of their range are occupied by different subspecies. In the past, morphologically diverse populations of Syncerus caffer have been associated with at least 92 scientific names, many of which were considered species, not subspecies. Today we consider the species polytypic, with three to four subspecies commonly recognized.

Syncerus caffer caffer, classic Cape or savannah buffalo, is found in the east and south, starting in southwest Ethiopia and through Kenya, Uganda, Rwanda, Tanzania, Zambia, and Malawi. The distribution is patchier through Angola, Mozambique, and Swaziland. In South Africa they are well distributed, except in the southeast and southwest, where they are absent.

Syncerus caffer nanus, forest buffalo, is found in the rainforests of west and central Africa. Their distribution is split into two main groups. The first is located from south Guinea moving east through Sierra Leone, Liberia, south Ivory Coast and southwest Ghana. The second distribution begins just to the west of the Nigeria-Cameroon border in south Cameroon, encompassing the area south along the coast including the Republic of the Congo and east, the north half of the Democratic Republic of the Congo and a piece of southwest Central African Republic. A contact zone occurs with Syncerus caffer caffer on the west end, following the border of the Democratic Republic of Congo.

Syncerus caffer brachyceros is the West Africa savannah buffalo. Its distribution lies just north of Syncerus caffer nanus, starting at south Senegal and north Guinea and continuing through southwest Mali, north Ivory Coast, Ghana, and Benin. Like Syncerus caffer nanus, the distribution is divided. The distribution begins at the Nigeria-Cameroon border and encompasses north Cameroon and the western edge of the Central African Republic. Along the southern edge of their border, they share a contact zone with Syncerus caffer nanus.

Syncerus caffer aequinoctialis, Central Africa savannah buffalo, share contact zones with all three other subspecies. On the east end, in east Central African Republic, their distribution briefly borders the distribution of Syncerus caffer brachyceros. The southern border of their distribution follows the northern border of the range for Syncerus caffer nanus. On the southwestern end in Ethiopia, the distribution meets the distribution of Syncerus caffer caffer. Their range encompasses most of Central African Republic, and portions of south Chad, South Sudan, and east Ethiopia. This subspecies is not always recognized. (Mloszewski, 1983; Sinclair, 1977; van Hoft and Prins, 2013)

Habitat

African buffalo can inhabit practically any habitat with permanent water and grass. They are found in dense lowland forests, lowland rainforests, montane forests and grasslands, Acacia grasslands, miombo (Brachystegia) woodlands, coastal savannas, plains and semi-arid bushland. They can be found to altitudes exceeding 4000 m. Buffalo show a preference for riverine habitat, especially in the dry season. They prefer close proximity (≤1 km) to water and are only found within 20 km of water. Buffalo density is correlated to annual precipitation, especially in areas without a large amount of permanent drinking water. Buffalo are rarely found where annual precipitation is less than 500 mm.

African buffalo are most associated with Acacia savannahs, miombo woodlands, montane woodlands, and the veld. Mixed-sex herds use open habitat more than bachelor herds. Syncerus caffer brachyceros are found in the Sahelian savannah and gallery forests. Syncerus caffer aequinoctialis are also found in the Sahelian savannah. Syncerus caffer nanus are found in rainforest regions with annual precipitation of at least 1500 mm. Specifically, they prefer rainforest clearings and open forest. (Hay, et al., 2008; Melletti, et al., 2007; Prins, 1996; Sinclair, 1974; Sinclair, 1977; Smitz, et al., 2013; van Hoft and Prins, 2013)

  • Range elevation
    4000 (high) m
    13123.36 (high) ft

Physical Description

African buffalo are very large, even-toed ungulates, characterized by their stocky build and heavy horns. Horns are present on both sexes and they are not ridged. Above their rounded hooves they have dewclaws. When identifying by skull, African buffalo can be distinguished from other bovids by short horn cones located posterior to the orbits and a distinctive fourth premolar with a fused paraconid and metaconid. Their skulls also have a relatively short facial region and lack preorbital fossae and ethmoid fissures. Many of these characteristics are variable. The dental formula is I0/3 C0/1 PM3/3 M3/3=32.

Sexual dimorphism exists in this species for size and a few physical characteristics. Males have a hump just past their thick necks and can have hairy skin-folds and a dewlap under their chins. Depending on the subspecies, a heavy, obvious boss is found on the top of the horns of males, which is supported by a bony ridge above the eyes. Horns of females are relatively slender and lack the boss. African buffalo exhibit variable sizes based on both sex and subspecies. In the largest subspecies, Syncerus caffer caffer, males can weight over 835 kg while females weigh up to 500 kg. Individuals in this subspecies stand 140 to 160 cm at the shoulder. The horn span is wide and can reach 130 cm. By comparison, Syncerus caffer nanus individuals stand only 105 cm at the shoulder and weighs under 250 to 320 kg. The boss on the horns of males in this subspecies is absent or heavily reduced. Horns are straighter, narrower in span and point backward. The two “transition zone” subspecies are intermediate in size. Syncerus caffer brachyceros individuals are smaller than Syncerus caffer caffer at 400 to 700 kg and their horns somewhat resemble those of domestic cattle, with bosses less developed than in Sycerus caffer caffer but more obvious than in Syncerus caffer nanus. Syncerus caffer aequinoctialis individuals are also 400 to 700 kg, with horns intermediate between S. c. caffer and S. c. brachyceros. There is a relationship between size and abundance of grass in the area, as the largest subspecies, Syncerus caffer caffer, is found where grasses are most plentiful while the smallest, S. c. nanus, is found in forests where grass is scarce. The transition-zone subspecies live in areas of moderate grass availability.

Color dimorphism exists in African buffalo and is loosely associated with subspecies. Syncerus caffer caffer individuals are typically dark brown to black. Syncerus caffer nanus individuals are typically red, but can also be black. The transition-zone subspecies are intermediate, and are often a sooty brown, possibly with a red tinge. Color often changes as a newborn buffalo matures and typically darkens with age until maturity. In general, African buffalo show considerable variation in size, color and horn shape within each subspecies, and even within herds.

Immature African buffalo can be aged by size, horn shape, and teeth eruption. Inconsistencies exist in the literature about the exact age when teeth erupt, but there are general patterns. Temporary teeth are in place by three to four months and have the dental formula i0/3 c0/1 pm3/3 = 20. The following sequence of eruptions apply to the lower jaw. By nine months the first permanent molar should be in the process of erupting, if not in use. By one and half years to one year and nine months, the second molar usually erupts. By two years nine months the third molar should be erupting and the first incisor is in use. By three and a half years to three years and eight months the second incisor and the second premolar should be in use, and the third premolar erupting. Between four and a half years and four years eight months all permanent teeth but the canine tooth should be in use. The last tooth to erupt is the incisor-like canine, which appears between four and a half and five and a half years of age. Teeth from the upper jaw experience a delay in eruption by one to three months. Buffaloes older than five can be aged with crown height measurement and cementum line counts.

It takes females four years and males five years to complete horn development. Until two years old the sexes look alike. At birth buffalo have small stubs of horns, which grow straight for the first 6 to 9 months. By their first birthday the horns have developed a curve. As young males grow, their horns thicken at the base to eventually join, forming the unified boss at the top of their head. Skin and hair is gradually lost over the boss. By two years of age the horn is halfway to meeting in the center and the bony ridge developing to support the developing boss becomes visible. Hair loss begins at this age. By age three to five, sub-adult males have a mostly completed boss that still has some hair and skin in the center. At this age the horn still has a pale outer layer. The boss is completed and the outer layer is gone around 5 years of age, but young males still lack the thick neck typical of older males. These males have few scars. Middle-aged males, between eight and ten, display ridges on the boss of their horns and have developed the typical thick necks. Males older than ten years show wear and possibly breakage on their horns, and heavy scarring and hair loss over their body and face.

Hybridization occurs between subspecies and intermediates can be found at contact zones. There has also been at least one hybridization between S. c. caffer and S. c. nanus in captivity. This pairing is notable because the two subspecies have different numbers of chromosomes, at 52 and 54 respectively. Some studies on genetic lineages support two subspecies: S. c. caffer and S. c. nanus, the latter of which would include S. c. brachyceros and S. c. aequinoctialis. Genetic differentiation between the three Central and West Africa subspecies is low, though these buffalo display higher genetic distance than that seen in S. c. caffer. Central and West Africa buffalo also have the highest number of private alleles. In general, S. c. caffer have very high within-population genetic variation and low between-population variation. They display a significant relationship between geographical distance and genetic distance. (Cribiu and Popescu, 1980; Grimsdell, 1973; Mloszewski, 1983; Sinclair, 1977; Skinner and Chimimba, 2005; Smitz, et al., 2013; Taylor, 1988; van Hoft and Prins, 2013; van Hooft, et al., 2000; van Hooft, et al., 2002)

  • Sexual Dimorphism
  • male larger
  • Range mass
    < 300 to > 835 kg
    to lb

Reproduction

African buffalo are promiscuous. As the mating season nears, males in bachelor herds rejoin mixed herds to begin seeking proestrous females. During their time in bachelor herds, males improve their body condition in preparation for mating. Upon finding a proestrous female, a male will “tend” her by following her around until she is receptive to mating. If approached by a more dominant male, tending males readily return to grazing and the new male takes over. Once the female is receptive, the tending male will mount and copulate at least twice within a half hour. During this time other males gather around them. When the mating male is finished, another male will tend the female. There is no permanent social bond between males and females. (Mloszewski, 1983; Sinclair, 1977)

Mating and births can occur throughout the year because female buffalo are polyestrus, with a cycle lasting 21 to 22 days. Estrus itself typically lasts 24 hours or less. A “breeding season” can generalized based on mating peaks, which are usually towards the end of the wet season, as the majority of births occur during the wet season about 340 days later. The breeding season is correlated with physiological changes in males, including increased ejaculate and sperm motility, as well as greater variation in circulating concentrations of testosterone. Gestation length is longer than expected for their size. It is thought to be protracted so that mothers experience optimal conditions both at conception and at parturition. Synchronized births are a common anti-predator adaptation in ungulates. The inter-birth period for African buffalo is variable, but can range from a year to over two years depending on population density, with an average of 18 to 19 months. Births typically occur when the herd is at rest, often in the early morning or afternoon. In captivity, newborns weighed between 24 and 60 kg, with a mean of 40 kg. Larger cows gave birth to heavier offspring, and twins are very rare. Time to weaning also depends on population density and can range from 4 months if the mother is killed or the calf is abandoned, to over a year and a half. Calves are typically weaned by 9 or 10 months, and begin eating solid foods by their second month. They become independent between one and two years of age. Females reach puberty around 3.5 years and are sexually mature (are likely to have conceived) by around 5 years. Males reach puberty by 2 to 3 years and are sexually mature by 4 to 6 years.

Males in the mixed herds constantly monitor females for signs of estrus. As the urine of females contains olfactory chemicals that signal bulls that they are approaching estrus. Males induce urination by licking the vulvae of females. After smelling the urine, males display the flehmen posture, with their chin raised, head thrust out, nostrils flared and upper lip curled showing upper gums. As a bull tends a female nearing estrus, he will occasionally test her willingness to mate by licking her and resting or rubbing his head on her rump. If she tolerates this behavior instead of moving away, the male will attempt to copulate. Receptive females may solicit copulation in the same way, by resting or rubbing her head on the bull, or by lowering her head under his belly. These behaviors, including mounting, also occur between members of the same sex.

Previous studies reported an even sex ratio among newborn calves, but this was often concluded after averaging data from one or more years. Recent research suggests that the sex ratio can vary with precipitation at conception, both seasonally and annually. Wet periods are associated with more males and dry seasons with more females. (Brown, et al., 1991; Mloszewski, 1983; Prins, 1996; Ryan, et al., 2006; Sinclair, 1977; Skinner, et al., 2006; Vidler, et al., 1963; van Hoft and Prins, 2013; van Hooft, et al., 2010)

  • Breeding interval
    The inter-birth period for African buffalo is between one year and over two years, depending on population density.
  • Breeding season
    Although mating happens throughout the year, a peak occurs at the end of the wet season.
  • Range number of offspring
    1 to 2
  • Average number of offspring
    1
  • Average number of offspring
    1
    AnAge
  • Range gestation period
    330 to 346 days
  • Average gestation period
    340 days
  • Range weaning age
    4 to >18 months
  • Average weaning age
    9-10 months
  • Range time to independence
    1 to 2 years
  • Range age at sexual or reproductive maturity (female)
    3.5 to 5.5 years
  • Average age at sexual or reproductive maturity (female)
    5 years
  • Range age at sexual or reproductive maturity (male)
    6 (high) years
  • Average age at sexual or reproductive maturity (male)
    4.5 years

African buffalo females provide extensive care for their young. Males do not invest in offspring. A female gives birth while still with the mixed herd, but the female is left behind with her newborn calf by the herd as they move on to forage. Newborns are less precocial than many other ungulates, such as wildebeest. Females wait with newborns as they gain strength, sometimes taking shelter in dense thickets during the day. They may leave the thicket to engage threats, such as charging at cars, then return to their calf. They rarely abandon calves to rejoin their herd. The calf is strong enough to rejoin herd with mother later in the day, though it can only walk slowly and must rest frequently. Calves cannot run well for the first several weeks, and fall behind as their mothers stay with the herd. Often a lost calf can reunite with its mother by catching up after the herd has slowed down, but if not, a lost calf will vocalize and actively seek its own mother. Some sources report mothers frequently leaving their calves behind and showing little anxiety or effort to find them beyond a certain distance, while other sources report the mother “checking in” on its calf by returning to it during flight.

Beyond feeding, defending, and guiding, maternal care consists of stroking, huddling, and playing. Play in buffalo calves is expressed by running, prancing and head-butting their mothers. Calves begin to learn to graze and drink water at two months. They remain largely dependent on a milk diet until four months of age, at which point they begin to rely less on milk.

Non-relatives in the herd may provide care for lost, orphaned, or abandoned calves. Solitary lost calves are sometimes accompanied by an older buffalo until it finds its mother. Abandoned or orphaned calves between 4 to 6 months may try to associate with another female, who sometimes accepts them. These “nanny” females are typically older females. They may adopt multiple calves, if more than one is orphaned, and will watch over them maternally. Calves in turn follow the lead of the adoptive female. Rarely adult bulls may initiate play with juveniles. (Mloszewski, 1983; Ryan, et al., 2006; Sinclair, 1977)

  • Parental Investment
  • precocial
  • female parental care
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
  • pre-weaning/fledging
    • provisioning
      • female

Lifespan/Longevity

African buffalo can live up to 22 years in the wild and have been recorded living 29.5 years in captivity. Adult males experience a higher rate of mortality than adult females or sub-adults of either sex. In one study, the average age of death was approximately 11 years old for both sexes. Disease is the most common cause of mortality, followed by lion predation. Male buffalo are preyed on by lions at a higher average rate than females. Females are senescent at 15 years old, at which point their survival rate is 25% lower than females in their prime. (Jolles, 2007; Jones, 1993; Prins, 1996)

Behavior

Africa buffalo are social and congregate in herds ranging from a few individuals to over a thousand. They form two types of herds: large, mixed-sex and mixed-age herds, sometimes called “breeding herds,” and small, all-male “bachelor herds.” The size of herds is variable based on subspecies and location, with S. c. caffer herds ranging from an average of 20 in montane areas to over 1500 in the Serengeti area. It is possible some of the smaller groups are part of larger herds that have split apart. In general, herds are smaller in denser vegetation. Syncerus caffer nanus herds range from under 10 individuals to over 20, with a mean of approximately 12. Transitional buffalo herds average around 20, but herds nearer to the grasslands can have over 100 individuals. Variation exists among different populations. For example whether they demonstrate a fusion-fission society or migratory behavior. Large herds frequently split up and exhibit a fusion-fission structure, smaller herds do not. Most herds do not migrate, especially herds that would face difficulty migrating naturally due to habitat fragmentation, fences, or other barriers. Migrations of over 80 km have been recorded, with African buffalo exhibiting different ranges for the two seasons. They are not strictly diurnal, but exhibit activity throughout the 24 hours of a day, usually with rest periods of low activity in the early morning and late afternoon. Mixed herds are mostly composed of adult females with their young and sub-adults (3 to 5 years old), and a few transient adult males. Adults make up 72% of the population, sub-adults 22%, and young approximately 6%. Outside of the breeding season, the ratio of adult males to females is less than 1:100, but the overall sex ratio for adults in a population is about 1:5. Including all ages, it isless than 1:2. During the breeding season the ratio in the mixed herd becomes 1:4. Mixed herds are sometimes described as broken down into subgroups. Sub-adult and two-year-old males form groups while staying within the mixed herd and sub-adult females stay with their mothers’ subgroups. Bachelor herds are smaller and consist solely of males, especially older males. Sub-adult males are rarely found in bachelor herds. Unlike adult females, males from bachelor herds regularly go back and forth among these two types of herds. The size of bachelor herds is seasonally variable, with adult males least likely to be in bachelor herds during the breeding season. Sub-adult males in bachelor herds do not show this seasonal variation in herd movement. Bachelor and mixed herds exhibit some habitat segregation, and bachelor herds travel less than mixed herds, resulting in males in bachelor herds gaining more body mass than males in mixed herds. Males have a dominance hierarchy in both mixed and bachelor herds. Despite herds containing potentially hundreds of members, males seem to always be aware of where their status in the dominance hierarchy. Dominance is most likely based on the body condition difference between the two interacting males, though it has also been speculated that a component of this hierarchy is endocrinal in nature. Males may spar to determine dominance. Frequency of sparring appears to be different among populations. Sparring is initiated when one male approaches and presents his horns to another male, who responds similarly. They lock horns and twist them from side to side. In adult bulls, the average is seven bouts of this behavior, lasting 10 seconds each per sparring session. After sparring, males typically return to grazing, as sparring almost never escalates beyond what appears to be mild competition. Dominance interactions are rare and typically include threat displays. Occasionally males fight by colliding once head-on against the boss of their horns. The winner is immediately decided by speed and weight and the loser is often chased a short distance. More serious fights are extremely rare and can end in death of one or both of the combatants. Herds seem to exhibit communal decision making when it comes to choosing where to travel. During the morning and afternoon rest periods, when the herd is mostly lying on the ground, a few adult cows will sequentially stand up and gaze off into the same direction. Later, when the herd is rousing, the first individuals to start moving all travel in that direction towards their new grazing location. Others follow their lead. Decisions about where to graze seem to be determined by the females, because if the males in the front stop to graze before their predetermined destination has been reached, the females behind simply continue without them. An interesting behavior observed in African buffaloes is seemingly altruistic partnerships. In one case, a partnership between two old males expressed itself as the healthier of the two assisting his blind and ailing companion. The more able bull would signal to the other when and in what direction to move and when to stop. (Brown, et al., 1991; Hay, et al., 2008; Korte, 2008; Megaze, et al., 2012; Melletti, et al., 2007; Mloszewski, 1983; Naidoo, et al., 2012; Naidoo, et al., 2014; Prins and Iason, 1989; Prins, 1996; Ryan, et al., 2006; Sinclair, 1977; Turner, et al., 2005)

Home Range

Home ranges can range from under 50 square kilometres to over 1000 square kilometres, depending on factors like amount of grass and water, density, and interspecific competition. It is not uncommon for home ranges to overlap and herds will sometimes briefly merge if they encounter one another in the overlap. Herds often follow familiar routes through their home ranges. (Grimsdell, 1969; Mloszewski, 1983)

Communication and Perception

African buffalo have acute senses to detect threats such as predators. They rely on a combination of visual, olfactory, and auditory cues to navigate their environment, especially for their anti-predatory behaviors. They have keen eyesight and are capable of spotting lions from over a kilometer away. Most communication is visual and takes the form of body language, for example initiating sparring involves the lowering of horns. African buffalo watch conspecifics for signs of threat and pay specific attention to individuals in alert postures. Communal “voting” on where to graze next is also done silently and visually, with buffaloes noting the direction of the initiating individuals face. African buffalo also rely on olfactory cues. Adult bulls use olfactory information in the urine of females to determine her stage in the estrous cycle. African buffalo pathfinders will break away from the herd and use a combination of sniffing the air and listening to the surrounding sounds before advancing. Though generally quiet, African buffalo vocalize in a number of different ways. Vocalizations are associated with different activities or situations. A notable example of a vocalization is the bleating of a calf in distress, which occurs when it becomes separated from its mother. This noise draws attention from mothers in general and other older individuals in the vicinity. Artificially reproducing this noise prompted responses by individuals and the herd, with the herd gathering around and individual females replying. (Grimsdell, 1969; Mloszewski, 1983; Sinclair, 1977)

Food Habits

African buffalo are grazing ruminants and devote a large proportion of their time to feeding and extracting nutrients from food. After grazing grass and sedges, they must spend time chewing their cud, or bolus, to extract more nutrients from their food. They prefer the leaves of grass, which dominate their diet during the wet season. Generally African buffalo prefer grasses with a higher leaf to stem ratio, such as from the genus Themeda. These grasses were preferentially grazed during an experiment with domestic African buffalo. They are pure or mixed grazers, with C4 plants making up the bulk of their diet, varying geographically from being entirely C4 plants in some areas to containing around 10% C3 plants in their diet in other areas. This variance in plant type is from supplemental browsing, especially in S. c. nanus. African buffalo typically show a pattern of activity marked by grazing, followed by ruminating, followed by rest. Activity patterns vary seasonally and by other factors such as predation pressure. For example, the grazing period in one study increased from an average of 1.5 hours in July to 4.5 hours in April. Resting periods show a similar variation. The rate at which the bolus is chewed shows a decline with age. (Cerling, et al., 2003; Codron, et al., 2007; Prins, 1996; Sinclair and Gwynne, 1972; Sinclair, 1977)

  • Plant Foods
  • leaves

Predation

African buffalo are formidable prey because of their large size and their aggression. They display a variety of anti-predator adaptations, the most obvious one being their congregation in large herds to increase vigilance and decrease the likelihood that any individual may be singled out. They are also willing to initiate or reciprocate aggression. The six main categories of buffalo anti-predator behavior are heightened caution, scrutiny, auditory-olfactory examination, flight, preventive aggression, and reciprocal aggression. Flight and reciprocal aggression are the most common high-intensity reactions to threat, especially in populations that are not under heavy predation pressure. Heightened caution manifests as alert behavior, with African buffalo holding their heads high and scanning. While displaying heightened caution, they may choose to detour around dangerous areas or move on from their current location. African buffalo attempt to locate a potential threat, usually visually, by scrutinizing their environment. To do this, they remain stationary and adjust the position of their head depending on distance and angle of the potential threat. While traveling through areas associated with danger, individuals stop, orient themselves to face backwards along their route, and scrutinize the area for up to 80 seconds. If a threat is detected the individual stands at alert, usually quickly steps forward, then stands alert again before turning around to run in the original direction, initiating a stampede. This sequence of actions, before the stampede, causes individuals in the vicinity, or the entire herd post-stampede, to be alert to that individual. In hilly terrain they preferentially watch from the tops of hills. Auditory-olfactory examination is carried out by a pathfinder leading the herd in an unfamiliar area. The individual stands apart from the herd and stands still for up to eight minutes, listening and smelling the air. Afterwards the pathfinder chooses to lead the herd forward, in a different direction, or remains stationary, depending on whether a threat was detected. Flight is the most common anti-predator response to serious threat, especially when flight is likely to be successful. Typically defensive aggression is initiated if flight is prevented, and aggression in general is intended to increase the likelihood of successful flight. Stampedes may or may not involve the entire herd, especially if only a portion of the herd faces threat. Stampedes begin with one or a few individuals and spread as those nearby join the stampede. Stampedes last varying distances depending on how clear the threat appears. For a confirmed serious threat, stampedes can last up to a kilometer, or until young buffalo are too tired to continue. After the end of the stampede, the herd typically continues on its route but in a roundabout way. Otherwise, stampedes stop a short distance away and the initiators reevaluate and either continue fleeing forward, go in a different direction, or resume their original movement. Lions (Panthera leo) are the primary predators of buffalo over a year in age. African buffalo can outrun lions by about 10 km per hr so, to be successful, lions must ambush them from a close distance. Depending on the region, lions either attack African buffalo by chance encounter or target them specifically. When they are not targeted, lions tend to attack individuals on the outskirts of the herd. They are likelier to attack males because males more than females tend to be at the periphery of a main herd or in separate bachelor herds. Males in bachelor groups are four times likelier to be killed by lions than females in mixed herds. Calf predation does not seem to be significant, possibly due to the protection of young by adults. In areas where lions specialize on preying on African buffalo, they tend to ambush them along a travel route. African buffalo in these areas have adapted to this predation pressure by anticipating where an ambush is likely to occur and deliberately protecting breeding females and the young by pushing males and invalids to the periphery. African buffalo that experience high predation pressure from lions are also likely to demonstrate more aggressive behavior toward them. Adult males are more aggressive and more successful at defending against lions than adult females. Bulls have been recorded to successfully fend off more than one lion, though this usually results in severe injuries. After throwing off attacking lions, bulls may display their horns to them and slowly retreat until the threat is over. African buffalo may also preemptively act aggressively toward non-threatening lions. Males have been known to approach lions from one km away to chase them. They sometimes chase lions into trees and will kill lions that fail to escape. Leopards (Panthera pardus) prey on buffalo calves. Reports of their significance as a predator are mixed. In some areas leopards do not seem to prey on African buffalo or are a negligible threat. One study in the forests of Lopé National Park in central Gabon found that forest buffalo (S. c. nanus) formed 13% of the diet of leopards. Spotted hyenas (Crocuta crocuta) and African wild dogs (Lycaon pictus) are minor predators of African buffalo. Spotted hyenas are not often observed attacking buffalo, but scat analyses show they make up a large portion of their diet in some studies. This can likely be explained as a result of them scavenging carcasses, often from lion kills. Because African buffalo herds can easily fend off hyenas packs, attacking one is a risky endeavor for hyenas. Reviews of hyena prey preference show that they avoid African buffalo. African wild dogs also strongly avoid African buffalo. They rarely attack them and only attack calves or juveniles. (Cronje, et al., 2002; Hay, et al., 2008; Hayward, et al., 2006a; Hayward, et al., 2006b; Hayward, 2006; Henschel and Skinner, 1990; Henschel, et al., 2005; Mloszewski, 1983; Prins and Iason, 1989; Prins, 1996; Sinclair, 1977; Wentworth, et al., 2011; di Silvestre, et al., 2000)

Ecosystem Roles

African buffalo are important grazers in the habitats they occupy. They cyclically return to graze patches, returning when the new growth is sufficiently high to be grazed again. In this way they manipulate the vegetation of their habitat. Their grazing opens the vegetation for other species that graze with the herd or follow in the wake of a buffalo herd. Species found in association with African buffalo in large herds include other ungulates, especially zebras and wildebeest. These species may also compete with buffalo for forage when food is scarce. Other ungulates may benefit from possible protection against lions by associating with the more aggressive African buffalo. African elephants are potentially a major competitor of African buffalo and agonistic interactions between the two species are common, with elephants initiating the aggressive encounter 90% of the time. African buffalo are host to a large variety of parasites, both internal and external. Examples include ticks, helminthes, trematodes, cestodes, nematodes, and pentastomids. Ticks are common vectors for diseases. Theileria, responsible for corridor disease, are hosted by Rhipicephalus appendiculatus. Ehrlichia ruminantium (previously Cowdria ruminantium) causes heartworm and are hosted by Amblyomma hebraeum and Amblyomma variegatum. These diseases can be transmitted to domestic cattle. African buffalo have a mutualistic relationship with red-billed oxpeckers and yellow-billed oxpeckers, who pick off ticks and other external parasites from large ungulates. Some research suggests that this relationship is not appreciated by the ungulate hosts, who attempt to displace them. African buffalo show somewhat less avoidance behaviors than other ungulates but still resist to some degree. This relationship may therefore be more parasitic in nature. Also associated with African buffalo are cattle egrets, which feed on insects disturbed by the them as they graze. African buffalo may charge at cattle egrets in their proximity. (Allsopp, et al., 1999; Bishop and Bishop, 2013; Dale, 1992; Grimsdell, 1969; Mloszewski, 1983; Norval, et al., 1994; Peter, et al., 2002; Prins, 1996; Sinclair, 1977; de Boer and Prins, 1990)

  • Ecosystem Impact
  • creates habitat
Commensal/Parasitic Species

Economic Importance for Humans: Positive

African buffalo are economically important for both the tourism and the trophy hunting industries. Commercial uses of wildlife on private land, such as safari hunting, wildlife harvesting (for meat and skins), and ecotourism provide lucrative income for farmers in regions where this type of private land use is permitted. Private land holdings with populations of African buffalo can charge up to twice as much as for ecotourism and hunting as competitors, providing incentive to have buffalo on the land. These ranches and private reserves also provide employment for local communities. African buffalo are illegally hunted or poached for bushmeat. (Carruthers, 2008; Gandy and Reilly, 2004; Lindsey, et al., 2013; Lindsey, et al., 2009; Sinclair, 1977; Williams, et al., 2000)

  • Positive Impacts
  • food
  • body parts are source of valuable material
  • ecotourism

Economic Importance for Humans: Negative

African buffalo are considered crop pests and disease vectors by farmers and ranchers. Like wild ungulates worldwide, they may be a nuisance in agricultural fields. Buffalo are also blamed for transmitting disease to domestic cattle herds. Buffalo can act as a reservoir for bovine tuberculosis (Mycobacterium bovis), re-infecting cattle herds or spreading the disease to healthy herds. A growing concern is foot-and-mouth disease and the potential for transmission. Risks of foot-and-mouth disease to humans is increased due to unsafe food handling practices, the consumption of organ meats, and a high prevalence of HIV/AIDS in the population, which reduces immune response. Another possible threat is brucellosis (g. Brucella) infection via the consumption of infected bushmeat. African buffalo are dangerous animals because of their size, aggressive nature, and formidable horns. (Alexander, et al., 2012; Clifford, et al., 2013; Michel and Bengis, 2012; Sinclair, 1977)

Conservation Status

All subspecies are considered together by IUCN and are listed as Least Concern. In the 1800's African buffalo populations were exposed to rinderpest, a disease of domestic cattle. In the following century rinderpest decimated buffalo populations. While populations have mostly recovered, new threats include habitat fragmentation and poaching. (Mloszewski, 1983; Sinclair, 1977; van Hoft and Prins, 2013)

Contributors

Christoph Ng (author), University of Manitoba, Jane Waterman (editor), University of Manitoba, Tanya Dewey (editor), University of Michigan-Ann Arbor.

Glossary

Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

World Map

acoustic

uses sound to communicate

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

causes or carries domestic animal disease

either directly causes, or indirectly transmits, a disease to a domestic animal

chemical

uses smells or other chemicals to communicate

colonial

used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.

dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

ecotourism

humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

folivore

an animal that mainly eats leaves.

food

A substance that provides both nutrients and energy to a living thing.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

herbivore

An animal that eats mainly plants or parts of plants.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

migratory

makes seasonal movements between breeding and wintering grounds

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

riparian

Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).

scrub forest

scrub forests develop in areas that experience dry seasons.

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

tactile

uses touch to communicate

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

year-round breeding

breeding takes place throughout the year

young precocial

young are relatively well-developed when born

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