View source for African Buffalo

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{{speciesbox
| status = LC|status_system = IUCN3.1
| trend = down
| status_ref = <ref name=iucn>IUCN SSC Antelope Specialist Group 2008. [http://www.iucnredlist.org/details/21251/0 ''Syncerus caffer''.] The IUCN Red List of Threatened Species. Version 2014.3. Downloaded on 08 April 2015.</ref>
| name = African buffalo
| fossil_range = {{Fossil range|0.7|0}}<small>Middle Pleistocene-Holocene</small>
| image = Serengeti Bueffel1.jpg
| image_caption = An African buffalo at Ngorongoro Conservation Area, Tanzania.
| taxon = Syncerus caffer
| parent_authority = Brian Houghton Hodgson, 1847
| authority = (Anders Sparrman, 1779)
| subdivision_ranks = Subspecies
| subdivision = 
''S. c. caffer''<br/>
''[[African Forest Buffalo|S. c. nanus]]''<br/>
''S. c. brachyceros''<br/>
''S. c. aequinoctialis''<br/>
''S. c. mathewsi''
}}
The '''African buffalo''' or '''Cape buffalo''' (''Syncerus caffer'') is a large African bovine.<ref name="MSW3">{{MSW3 Artiodactyla | id = 14200707 | pages = 695-696}}</ref> It is not closely related to the slightly larger wild water buffalo of Asia and its ancestry remains unclear. ''Syncerus caffer caffer'', the Cape buffalo, is the typical subspecies, and the largest one, found in South and East Africa. ''S. c. nanus'' (African forest buffalo) is the smallest subspecies, common in forest areas of Central and West Africa, while  ''S. c. brachyceros'' is in West Africa and ''S. c. aequinoctialis'' is in the savannas of Central Africa. The adult buffalo's horns are its characteristic feature; they have fused bases, forming a continuous bone shield referred to as a "boss".  They are widely regarded as very dangerous animals, as they gore and kill over 200 people every year.

The African buffalo is not an ancestor of domestic cattle and is only distantly related to other larger bovines. Owing to its unpredictable nature, which makes it highly dangerous to humans, the African buffalo has never been domesticated, unlike its Asian counterpart, the water buffalo. Other than humans, African Cape buffaloes have few predators aside from lions, and are capable of defending themselves. Being a member of the big five game, the Cape buffalo is a sought-after trophy in hunting.

==Description==
[[File:Syncerus caffer african buffalo skull MNHN.jpg|thumb|left|Skull of an African buffalo]]
The African buffalo is a very robust species. Its shoulder height can range from {{convert|1.0|to|1.7|m|ft|abbr=on}} and its head-and-body length can range from {{convert|1.7|to|3.4|m|ft|abbr=on}}. Compared with other large bovids, it has a long but stocky body (the body length can exceed the wild water buffalo, which is heavier and taller) and short but thickset legs,  resulting in a relatively short standing height. The tail can range from {{convert|70|to|110|cm|in|abbr=on}} long. Savannah-type buffaloes weigh {{convert|500|to|1000|kg|lb|abbr=on}}, with males normally larger than females, reaching the upper weight range.<ref>Raphael, Marcel (2006) [http://iml.jou.ufl.edu/projects/Fall06/Raphael/pages/African%20Buffalo.html African Buffalo].</ref>  In comparison, forest-type buffaloes, at {{convert|250|to|450|kg|lb|-2|abbr=on}}, are only half that size.<ref name="ultimateungulate">{{cite web|first=Brent |last=Huffman|url=http://www.ultimateungulate.com/Artiodactyla/Syncerus_caffer.html |title=Syncerus caffer – African buffalo |publisher=Ultimateungulate.com|date=2010-05-24 |accessdate=2010-10-23}}</ref><ref>Boitani, Luigi, ''Simon & Schuster's Guide to Mammals''. Simon & Schuster/Touchstone Books (1984), ISBN 978-0-671-42805-1</ref> Its head is carried low; its top is located below the backline. The front hooves of the buffalo are wider than the rear, which is associated with the need to support the weight of the front part of the body, which is heavier and more powerful than the back.

Savannah-type buffaloes have black or dark brown coats with age. Old bulls have whitish circles around their eyes. Females tend to have more-reddish coats. Forest-type buffaloes are reddish brown in colour with horns that curve back and slightly up. Calves of both types have red coats.

[[File:Syncerus range map.png|thumb|Range of the commonly accepted forms of the African buffalo]]

A characteristic feature of the horns of adult male African buffalo is fusion of their bases, forming a continuous bone shield referred to as a "boss".  From the base, the horns diverge downwards, then smoothly curve upwards and outwards. In large bulls, the distance between the ends of the horns can reach upwards of one metre. The horns form fully when the animal reaches the age of five or six years. In cows, the horns are, on average, 10–20% smaller, and the boss is less prominent. Forest buffalo horns are smaller than those of the savannah buffalo, usually measuring less than {{convert|40|cm|in}}, and are almost never fused.

==Subspecies==
* ''Syncerus caffer caffer'' (Cape buffalo or southern savanna buffalo) is the typical subspecies, and the largest one, with large males weighing up to {{convert|910|kg|lb|abbr=on}}. It is peculiar to South and East Africa.  Buffaloes of this subspecies living in the south of the continent, notably tall in size and ferocity, are the so-called Cape buffalo. Color of this subspecies is the darkest, almost black.
* ''S. c. nanus'' (forest buffalo, dwarf buffalo) is the smallest subspecies; the height at the withers is less than 120&nbsp;cm and average weight is about {{convert|270|kg|lb|abbr=on}}. The color is red, with darker patches on the head and shoulders, and in the ears, forming a brush.  The dwarf buffalo is common in forest areas of Central and West Africa.  This subspecies is so different from the standard model, some researchers consider it still a separate species, ''S.  nanus''. Hybrids between the typical subspecies and dwarf are not uncommon.
* ''S. c. brachyceros'' (Sudanese buffalo) is, in morphological terms, intermediate between those two subspecies.  It occurs in West Africa.  Its dimensions are relatively small, especially  compared to other buffalo found in Cameroon, which weigh half as much as the South African subspecies (bulls weighing {{convert|600|kg|lb|abbr=on}} are considered to be very large).
* ''S. c.  aequinoctialis'' ([[Nile buffalo]]) is confined to the savannas of Central Africa. It is similar to the Cape buffalo, but somewhat smaller, and its color is lighter. This subspecies is sometimes included in the Sudanese buffalo.<ref name="Groves & Leslie, 2011">C. P. Groves, D. M. Leslie Jr. (2011) ''Family Bovidae (Hollow-horned Ruminants).'' pp. 585–588. In: Wilson, D. E., Mittermeier, R. A., (Hrsg.). ''Handbook of the Mammals of the World. Volume 2: Hooved Mammals.'' Lynx Edicions, 2009. ISBN 978-84-96553-77-4</ref>
* ''S. c. mathewsi'' (mountain buffalo or Virunga buffalo) is not universally recognized.  It lives in mountainous areas of the Democratic Republic of the Congo, Rwanda, and Uganda.<ref>{{cite web|url=http://www.biolib.cz/en/taxon/id726113/|title=BioLib - Syncerus caffer matthewsi (Virunga Buffalo)|work=biolib.cz}}</ref>

==Ecology==
The African buffalo is one of the most successful grazers in Africa. It lives in swamps and floodplains, as well as mopane grasslands and forests of the major mountains of Africa. This buffalo prefers a habitat with dense cover, such as reeds and thickets, but can also be found in open woodland.<ref name ="Estes 1991">Estes, R. (1991) ''The Behavior Guide to African Mammals, Including Hoofed Mammals, Carnivores, Primates.'' Los Angeles, The University of California Press. pp. 195–200 ISBN 0520080858</ref> While not particularly demanding with regard to habitat, they require water daily, so depend on perennial sources of water. Like the [[plains zebra]], the buffalo can live on tall, coarse grasses. Herds of buffalo mow down grasses and make way for more selective grazers. When feeding, the buffalo makes use of its tongue and wide incisor row to eat grass more quickly than most other African herbivores. Buffaloes do not stay on trampled or depleted areas for long.

Other than humans, African Cape buffaloes have few predators and are capable of defending themselves against (and killing) lions.<ref>{{cite web|url = http://nature.ca/notebooks/english/capebuff.htm|title = Cape Buffalo|publisher = Canadian Museum of Nature|accessdate=2010-10-23 }}</ref> Lions do kill and eat buffalo regularly, and in some regions, the buffaloes are the lions' primary prey. It typically takes several lions to bring down a single adult buffalo, usually the entire pride joins the hunt; however, several incidents have been reported in which lone adult male lions have been able to successfully bring down adult animals. The average-sized crocodile  typically attacks only old solitary animals and young calves, though they can kill healthy adults. Also, the Nile crocodile is the only animal that typically takes down adult buffalo alone, as pride attack is the preferred method of lions when taking down such large prey.<ref name="ultimateungulate"/> The cheetah, leopard, and spotted hyena are a threat only to newborn calves, though spotted hyenas have been recorded killing full-grown bulls on rare occasions.<ref name="Hyena">{{cite book | last= Kruuk |first= Hans | title=The Spotted Hyena: A study of predation and social behaviour | year= 1979 | page= 335 | isbn= 978-0-226-45508-2 | publisher= University of Chicago Press }}</ref>

===Diseases===
The Cape buffalo is susceptible to many diseases, including bovine tuberculosis, corridor disease, and foot and mouth disease. As with many diseases, these problems remain dormant within a population as long as the health of the animals is good. These diseases do, however, restrict the legal movements of the animals and fencing infected areas from unaffected areas is enforced. Some wardens and game managers have managed to protect and breed "disease-free" herds which become very valuable because they can be transported. Most well-known are Lindsay Hunt's efforts to source uninfected animals from the Kruger National Park in South Africa. Some disease-free buffaloes in South Africa have been sold to breeders for close to US$130,000.
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==Social behavior==
[[File:Afrikanische Büffel (Syncerus caffer) 1.jpg|thumb|right|Buffalo herd]]
Herd size is highly variable. The core of the herds is made up of related females, and their offspring, in an almost linear dominance hierarchy.  The basic herds are surrounded by subherds of subordinate males, high-ranking males and females, and old or invalid animals. The young males keep their distance from the dominant bull, which is recognizable by the thickness of his horns.
During the dry season, males split from the herd and form bachelor groups.<ref name ="Turner 1995"/> Two types of bachelor herds occur: ones made of males aged four to seven years and those of males 12 years or older.<ref name= "Ryan 2006">{{cite journal |last=Ryan |first=S. J. |last2=Knetchtel |first2=Christiane U. |last3=Wayne |first3=M. |year=2006|title=Range and habitat Selection of African Buffalo in South Africa|doi=10.2193/0022-541X(2006)70[764:RAHSOA]2.0.CO;2|url=http://nature.berkeley.edu/getzlab/Reprints06/RyanKnechtelGetzJWM06.pdf|volume= 70|issue=3|pages=764–776|journal=Journal of Wildlife Management}}</ref> During the wet season, the younger bulls rejoin a herd to mate with the females. They stay with them throughout the season to protect the calves.<ref name ="Main 1990">{{cite journal |last=Main |first=M. B. |last2=Coblentz |first2=Bruce E. |year=1990|title=Sexual Segregation among Ungulate: A Critique|jstor=3782137|journal=Wildlife Society Bulletin|volume=18|issue=2|pages=204–210}}</ref> Some older bulls cease to rejoin the herd, as they can no longer compete with the younger, more aggressive males. Males have a linear dominance hierarchy based on age and size. Since a buffalo is safer when a herd is larger, dominant bulls may rely on subordinate bulls and sometimes tolerate their copulation.<ref name ="Turner 1995"/>
[[File:Syncerus caffer fight1.jpg|thumb|Bulls in position to spar]]

Adult bulls spar in play, dominance interactions, or actual fights. A bull approaches another, lowing, with his horns down, and waits for the other bull to do the same thing. When sparring, the bulls twist their horns from side to side.<ref name="Sinclair 1977">Sinclair, A. R. E. (1977) ''The African Buffalo''. Chicago, The University of Chicago Press.</ref> If the sparring is for play, the bull may rub his opponent's face and body during the sparring session. Actual fights are violent but rare and brief. Calves may also spar in play, but adult females rarely spar at all.

African buffaloes are notable for their apparent altruism. Females appear to exhibit some sort of "voting behavior". During resting time, the females stand up, shuffle around, and sit back down again. They sit in the direction they think they should move. After an hour of more shuffling, the females travel in the direction they decide. This decision is communal and not based on hierarchy or dominance.<ref>{{cite journal |last=Wilson |first=D. S. |year=1997|title=Altruism and Organism: Disentangling the Themes of Multilevel Selection Theory|jstor=2463504|pmid=18811309|journal=The American Naturalist|volume=150|pages= S122–S134|doi=10.1086/286053}}</ref> When chased by predators, a herd  sticks close together and makes it hard for the predators to pick off one member. Calves are gathered in the middle. A buffalo herd responds to the distress call of a captured member and tries to rescue it.<ref name="Sinclair 1977"/> A calf's distress call gets the attention of not only the mother, but also the herd. Buffaloes engage in [[mobbing behavior]] when fighting off predators.  They have been recorded killing a lion<ref>{{cite av media|url=https://www.youtube.com/watch?v=PJjcQBSPDaI|title=YouTube|work=youtube.com}}</ref> and chasing lions up trees and keeping them there for two hours, after the lions have killed a member of their group. Lion cubs can get trampled and killed. In one videotaped instance, known as the Battle at Kruger, a calf survived an attack by both lions and a crocodile after intervention of the herd. 

===Vocalizations===
African buffaloes make various vocalizations. Many calls are lower-pitched versions of those emitted by domestic cattle. They emit low-pitched, two- to four-second calls intermittently at three- to six-second intervals to signal the herd to move. To signal to the herd to change direction, leaders  emit "gritty", "creaking gate" sounds.<ref name ="Estes 1991"/> When moving to drinking places, some individuals make long "maaa" calls up to 20 times a minute. When being aggressive, they make explosive grunts that may last long or turn into a rumbling growl. Cows produce croaking calls when looking for their calves. Calves make a similar call of a higher pitch when in distress.<ref name ="Estes 1991"/> When threatened by predators, they make drawn-out "waaaa" calls. Dominant individuals make calls to announce their presence and location. A version of the same call, but more intense, is emitted as a warning to an encroaching inferior.<ref name ="Estes 1991"/> When grazing, they make various sounds, such as brief bellows, grunts, honks, and croaks.


===Reproduction===
[[File:Cape Buffalo calf.jpg|thumb|right|Cape buffalo and her calf]]
Buffaloes mate and give birth only during the rainy seasons. Birth peak takes place early in the season, while mating peaks later. A bull closely guards a cow that comes into heat, while keeping other bulls at bay.<ref name ="Estes 1991"/><ref name ="Turner 1995">{{cite journal |last=Turner |first=W. C. |last2=Jolles |first2=A. E.|last3=Owen-Smith |first3=N. |year=2005|title=Alternating Sexual Segregation During the Mating Season By Male African Buffalo (''Syncerus caffer'')|doi=10.1017/S095283690500748X |journal=Journal of Zoology |volume=267|issue=3|pages=291–299}}</ref> This is difficult, as cows are quite evasive and attract many males to the scene. By the time a cow is in full estrus, only the most dominant bull in the herd/subherd is there.<ref name ="Estes 1991"/>

Cows first calve at five years of age, after a gestation period of 11.5 months. Newborn calves remain hidden in vegetation for the first few weeks while being nursed occasionally by the mother before joining the main herd. Older calves are held in the centre of the herd for safety.
<ref>{{cite web|url =http://www.bbc.co.uk/nature/species/African_buffalo|title = African Buffalo|publisher = British Broadcasting Corporation |accessdate=2010-10-23| archiveurl= https://web.archive.org/web/20101027125404/http://www.bbc.co.uk/nature/species/African_Buffalo| archivedate= 27 October 2010| deadurl= no}}</ref> The maternal bond between mother and calf lasts longer than in most bovids. However, when a new calf is born, the bonding ends and the mother keeps her previous offspring at bay with horn jabs. Nevertheless, the yearling follows its mother for another year or so. Males leave their mothers when they are two years old and join the bachelor groups. Young calves, unusually for bovids, suckle from behind their mothers, pushing their heads between the mothers' legs.<ref name="Prins1996">{{cite book |last=Prins |first=H.H.T |title=Ecology and Behaviour of the African Buffalo: Social Inequality and Decision Making|url=https://books.google.com/books?id=5FUs3eujlFwC&pg=PA118|accessdate=31 July 2013|year=1996|publisher=Springer|isbn=978-0-412-72520-3|page=118}}</ref>

==Relationship with humans==
[[File:Ernest Hemingway poses with water buffalo, Africa, 1953.jpg|thumb|Ernest Hemingway poses with a Cape buffalo he shot in 1953.]]

===Status===
The current status of African Cape buffalo is dependent on the animal's value to both trophy hunters and tourists, paving the way for conservation efforts through antipoaching patrols, village crop damage payouts, and CAMPFIRE payback programs to local areas.

The buffalo is listed as least concern by the IUCN "as the species remains widespread, with a global population estimated at nearly 900,000 animals, of which more than three-quarters are in protected areas. While some populations (subspecies) are decreasing, others will remain unchanged in the long term if large, healthy populations continue to persist in a substantial number of national parks, equivalent reserves and hunting zones in southern and eastern Africa."<ref name=iucn/>

In the most recent and available census data at continental scale, the total estimated numbers of the three African buffalo savanna subspecies (''S. c. caffer, S. c. brachyceros, S. c. aequinoctialis'') are at 513,000 individuals.<ref>Melletti M. and Burton J. (Eds). 2014. Ecology, Evolution and Behaviour of Wild Cattle. Implications for Conservation. Cambridge University Press</ref>

In the past, numbers of African buffaloes suffered their most severe collapse during the great rinderpest epidemic of the 1890s, which, coupled with pleuro-pneumonia, caused mortalities as high as 95% among livestock and wild ungulates.<ref>{{cite journal |last=Winterbach |first=H. E. K. |year=1998|title=Research review: the status and distribution of Cape buffalo Syncerus caffer caffer in southern Africa|journal=South African Journal of Wildlife Research|volume= 28|issue=3|pages=82–88|url=http://www.sabinet.co.za/abstracts/wild/wild_v28_n3_a4.html}}</ref>

Being a member of the big-five game family, a term originally used to describe the five most dangerous animals to hunt, the Cape buffalo is a sought-after trophy, with some hunters paying over $10,000 for the opportunity to hunt one. The larger bulls are targeted for their trophy value, although in some areas, buffaloes are still hunted for meat.

=== Attacks ===
One of the big five, it is known as "the Black Death" or "widowmaker", and is widely regarded as a very dangerous animal, as it gores and kills over 200 people every year. Buffaloes are sometimes reported to kill more
people in Africa than any other animal, although the same claim is also made of hippos and crocodiles.<ref>{{cite web|url = http://www.on-the-matrix.com/africa/buffalo.asp|first=Bruce G. |last=Stumpf  |title = Africa on the Matrix: The Cape Buffalo |accessdate=2010-10-23}}</ref> Buffaloes are notorious among big-game hunters as very dangerous animals, with wounded animals reported to ambush and attack pursuers.<ref>{{cite web|url = http://www.safaribwana.com/ANIMALS/animpages/buffalo.htm|title = African Animals Hunting facts and tips – Buffalo Hunting |publisher=safariBwana newsletter |accessdate=2010-10-23}}</ref>

==References==
{{reflist}}

==Further reading==
* Melletti M. and Burton J. (Eds). 2014. ''Ecology, Evolution and Behaviour of Wild Cattle. Implications for Conservation'' (Cambridge University Press). http://www.cambridge.org/us/academic/subjects/life-sciences/animal-behaviour/ecology-evolution-and-behaviour-wild-cattle-implications-conservation
* ''Ecology and Behaviour of the African Buffalo – Social Inequality and Decision Making'' (Chapman & Hall Wildlife Ecology & Behaviour)
* Huffman, B. 2006. [http://www.ultimateungulate.com ''The ultimate ungulate page'']. ''UltimateUngulate.com''. Retrieved January 9, 2007.
* International Union for Conservation of Nature and Natural Resources (IUCN). (2006) [http://www.iucnredlist.org/search/details.php/21251/all ''Syncerus caffer''],
* Nowak, R.M. and Paradiso, J.L. 1983. ''Walker's Mammals of the World''. Baltimore, Maryland, USA: The Johns Hopkins University Press. ISBN 978-0-8018-2525-5

[[Category:Mammals of Zambia]]
[[Category:Wildlife of Zambia]]