Plains Zebra

Plains Zebra
Equus quagga boehmi in the Ngorongoro Crater in Tanzania
Conservation status
Least Concern (IUCN 3.1)[1]
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	[Perissodactyla]
Family:	Equidae
Genus:	Equus
Subgenus:	Hippotigris
Species:	E. quagga
Binomial name
Equus quagga Boddaert, 1785
Subspecies
E. q. quagga † E. q. burchellii E. q. boehmi E. q. borensis E. q. chapmani E. q. crawshayi
Plains Zebra range
Synonyms[1]
Equus burchelli (Gray, 1824) [orth. error] Equus burchellii Equus burchellii Schinz, 1845

The plains zebra (Equus quagga, formerly Equus burchelli), also known as the common zebra or Burchell's zebra, is the most common and geographically widespread species of zebra.^[2] It ranges from the south of Ethiopia through East Africa to as far south as Angola and eastern South Africa. The plains zebra remains common in game reserves, but is threatened by human activities such as hunting for its meat and hide, as well as competition with livestock and encroachment by farming on much of its habitat.

Taxonomy

The Plains zebra and perhaps the mountain zebra belong to the subgenus Hippotigris, but Grévy's zebra is the sole species of subgenus Dolichohippus. The latter resembles an ass, while the former two are more horse-like. All three belong to the genus Equus along with other living equids. Recent phylogenetic evidence suggests that Grevy's zebras (and perhaps also Mountain Zebras) are with asses and donkeys in a separate lineage from the Plains zebra.^[3] In areas where Plains zebras are sympatric with Grévy's zebras, it is not unusual to find them in the same herds^[4] and fertile hybrids occur.^[5] In captivity, Plains zebras have been crossed with mountain zebras. The hybrid foals lacked a dewlap and resembled the plains zebra apart from their larger ears and their hindquarters pattern.

Subspecies

Quagga (Equus quagga quagga)

Burchell's Zebra (Equus quagga burchellii) in Etosha National Park, Namibia

In 2004, C.P. Groves and C.H. Bell investigated the taxonomy of the zebra genus, Equus, subgenus Hippotigris. They published their research in Mammalian Biology. They revised the subspecies of the plains zebra Equus quagga. Six subspecies are now recognizable. The completely maneless Somali population may represent a seventh subspecies: Equus quagga isabella (Ziccardi, 1958). This subspecies may be valid, but at present there is no evidence that it is.

• Quagga, †Equus quagga quagga – Boddaert, 1785
• Burchell's zebra, Equus quagga burchellii – Gray, 1824
• Grant's zebra, Equus quagga boehmi – Matschie, 1892
• Selous' Zebra, Equus quagga borensis – Lönnberg, 1921
• Chapman's zebra, Equus quagga chapmani – Layard, 1865
• Crawshay's Zebra, Equus quagga crawshayi – De Winton, 1896

The quagga was originally classified as a separate species, Equus quagga, in 1778. Over the next 50 years or so, many other zebras were described by naturalists and explorers. Because of the great variation in coat patterns (no two zebras are alike), taxonomists were left with a great number of described "species", and no easy way to tell which of these were true species, which were subspecies, and which were simply natural variants. The quagga was the first extinct creature to have its DNA studied. Recent genetic research at the Smithsonian Institution has demonstrated that the quagga was in fact not a separate species at all, but diverged from the plains zebra, between 120,000 and 290,000 years ago, and suggests that it should be named Equus burchelli quagga. However, according to the rules of biological nomenclature, where there are two or more alternative names for a single species, the name first used takes priority. As the quagga was described about thirty years earlier than the Burchell's zebra, it appears that the correct terms are E. quagga quagga for the quagga and E. quagga burchelli for the plains zebra, unless "Equus burchelli" is officially declared to be a nomen conservandum.

The Burchell's zebra was thought to have been hunted to extinction. However Groves and Bell concluded in their 2004 publication that "the extinct true Burchell's zebra" is a phantom. Careful study of the original zebra populations in Zululand and Swaziland, and of skins harvested on game farms in Zululand and Natal, has revealed that a certain small proportion shows similarity to what now is regarded as typical "burchellii". The type localities of the subspecies Equus quagga burchellii and Equus quagga antiquorum (Damara zebra) are so close to each other that the two are in fact one, and that therefore the older of the two names should take precedence over the younger. They therefore say that the correct name for the southernmost subspecies must be burchellii not antiquorum. The subspecies Equus quagga burchellii still exists in KwaZulu-Natal and in Etosha.

Physical description

Variation in coat pattern in zebras. The patterns show progressively more black left to right and top to bottom.

The plains zebra is mid-sized, smaller on average than the other two zebra species, and thick bodied with relatively short legs. There is some variation in size, based on the animals' condition and subspecies. Adults of both sexes can stand from 1.1 to 1.47 m (3.6 to 4.8 ft) high at the shoulder, are 2 to 2.5 m (6.6 to 8.2 ft) long, not counting a 50 cm (20 in) tail, and weigh 175 to 387 kg (390 to 850 lb). Males may weigh 10% more than females.^[6][7]

Like all zebras, they are boldly striped in black and white, and no two individuals look exactly alike. They also have black or dark muzzles. The natal coat of a foal is brown and white. All have vertical stripes on the forepart of the body, which tend towards the horizontal on the hindquarters. The northern populations have narrower and more defined striping;^[8][9] southern populations have varied but lesser amounts of striping on the underparts, the legs and the hindquarters.^[8] Southern populations also have brown "shadow" stripes between the black and white coloring.^[8][9] These are absent or poorly expressed in northern zebras.^[8][9]

An experiment was done at the Knoxville Zoo where a zebra was shaved. The underlying skin was black, not the previously thought white. The first subspecies to be described, the now-extinct quagga, had plain brown hindquarters. (Technically, because the quagga was described first as E. quagga, the proper zoological name for the most common form of the plains zebra is E. quagga burchelli.) There have been various mutations of the zebra's pelage from mostly white to mostly black.^[10] Rare albino zebras have been recorded in the forests of Mount Kenya.^[11]

Ecology

Range and habitat

Plains zebra are highly dependent on water.

The Plains zebra's range stops short of the Sahara from southern Sudan and southern Ethiopia extending south along eastern Africa, as far as Zambia, Mozambique, and Malawi, before spreading into most southern African countries. They are regionally extinct in Burundi and Lesotho, and they may have lived in Algeria in the Neolithic Era.^[12]

Plains zebras generally live in treeless grasslands and savanna woodlands^[9] but can be found in a variety of habitats, both tropical and temperate. However they are generally absent from deserts, dense rainforests and permanent wetlands.^[9] Zebras also live in elevations from sea level to 4,300 m on Mount Kenya. They rely on rainfall for food and water, and go on great migrations to follow the rains. The zebras will migrate up to 700 miles (1,100 km) for food. Other grazers also migrate. Plains zebras are highly water-dependent^[4] and are usually found within 25–30 kilometers of a water source.

Diet and predation

Zebras grazing with wildebeest in the Ngorongoro Crater

In one study, the zebra's diet was estimated to be 92% grass, 5% herbs, and 2% shrubs.^[13] Unlike many of the large ungulates of Africa, the plains zebra does not require (but still prefers) short grass to graze. It eats a wide range of different grasses, preferring young, fresh growth where available, and also browses on leaves and shoots from time to time. In consequence, it ranges more widely than many other species, even into woodlands, and it is often the first grazing species to appear in a well-vegetated area.^[4] Zebras have a simple stomach and use hindgut fermentation (caeco-colic) which allows them to digest and assimilate larger amounts of forage during a 24-hour period.^[14] Thus, zebras are less selective in foraging, but they do spend much time eating. The zebra is a pioneer grazer and prepares the way for more specialized grazers like blue wildebeests and Thomson's gazelles^[4] who depend on shorter and more nutritional grasses.

The plains zebra's major predators are lions and spotted hyenas.^[8] Nile crocodiles are also great threats during river crossings. Wild dogs, cheetahs, and leopards also prey on zebras, although the threats they pose are generally minor. Hyenas, dogs, cheetahs, and leopards mostly attack zebra foals.

Interactions with other grazers

Plains zebra herds will mix and migrate together along with other species such as wildebeests. Wildebeests and zebras generally coexist peacefully and will alert each other to predators. However, aggressive interactions occasionally occur.^[15][16]

Behavior

Social structure

A zebra harem in Etosha National Park

The plains zebra is highly social and usually forms small family groups called harems, which consist of a single stallion, several mares, and their recent offspring. The adult membership of a harem is highly stable, typically remaining together for months to years. Groups of all male "bachelors" also exist. These are stable groups of 2-15 males with an age-based hierarchy lead by a young male.^[4] These males stay in their groups until they are ready to start a harem. The bachelors spend much time "rehearsing" their adult roles with play fights and greeting/challenge rituals.^[4] Multiple harems and bachelor groups come together to form herds. Plains zebras are unusual among harem-holding species in forming these groups.^[17] In addition, pairs of harems may create temporarily stable subgroups within a herd.^[17] Zebra interactions with those outside their core group may reflect a second tier of social relationships that critically shape the society of the plains zebra. Among harem-holding species, this has only been observed in primates like the gelada and the hamadryas baboon.^[17]

Stallions form harems by abducting young mares from their family harems.^[4][18] When a mare reaches sexual maturity, she will exhibit the estrous posture, which attracts nearby stallions,^[18] both bachelors and harem leaders. Her harem stallion (usually her father) will chase off or fight stallions attempting to abduct her. Even after a young mare is isolated from her natal harem, the fight over her continues until her estrous cycle is over, and it starts again with the next estrous cycle.^[19] It is rare that the mare's original abductor keeps her for long.^[19] When the mare finally ovulates, the male that impregnates her keeps her for good. Thus, the mare becomes a permanent member of a new harem.^[19][20]

Two zebras fighting

Mares exist in a hierarchy, with the alpha female being the first to mate with the stallion and being the one to lead the group. When new mares are added to the group, they are met with hostility by the other mares. Thus, the stallion must shield the new mares until the aggression subsides.^[4][20] The most recently added females rank lowest. Females that become unfit or weak may drop in their rank, though. The female memberships of a harem stay intact even if a new stallion takes over. Zebras strengthen their social bonds with grooming. Members of a harem nip and scrape along the neck, shoulders, and back with their teeth and lips. Mothers and foals groom the most often, followed by siblings. Grooming shows social status and eases aggressive behavior.^[4]

A stallion will defend his group from other males. When challenged, the stallion would issue a warning to the invader by rubbing nose or shoulder with him. If the warning is not heeded, a fight breaks out. Zebra fights often become very violent, with the animals biting at each other's necks, heads or legs, wrestling to the ground, and occasional kicking. Sometimes a stallion will lie still on the ground as if surrendering, but once the other male lets up, will strike and continue the fight.^[4] Most fighting occurs over young mares in estrus, and as long as a harem stallion is healthy, he will usually not be challenged. Only unhealthy stallions have their harems taken over, and even then the new stallion gradually takes over pushing the old one out without a fight.^[4]

Communication

Zebras communicate with each other with high pitched barks and whinnying. A zebra's ears signify its mood. When a zebra is in a calm, tense or friendly mood, its ears stand erect. When it is frightened, its ears are pushed forward. When angry, the ears are pulled backward. When surveying an area for predators, zebras will stand in an alert posture, with ears erect, head held high, and staring. When tense, they will also snort. When a predator is spotted or sensed, a zebra will bark loudly.

Reproduction

Mother zebra nursing her foal

The stallion mates with all his mares. Mares may give birth to one foal every twelve months. The birthing peak is during the rainy season. She nurses the foal for up to a year. The stallion is generally intolerant of foals that are not his. It is possible that zebras practice infanticide and feticide, although such incidences have only been observed in captive individuals.^[21] Like horses, zebras are able to stand, walk, and suckle shortly after they are born. At the moment of birth, a mother zebra keeps any other zebra away from her foal, including the stallion, the other mares, and even the previous offspring. Later, though, they all bond. Plains zebra foals are protected by their mother as well as the head stallion and the other mares in their group. Even with parental protection, up to 50% of zebra foals are taken by predation, disease, and starvation each year.

Young male zebras eventually leave their family groups. This is not because of sexual maturity or being kicked out by their fathers, but because their relationship with their mothers have faded after the birth of a sibling.^[8][18] The young stallion then seeks out other young stallions for company.^[18] Young females may stay in the harem until they are abducted by another stallion.^[8]

Anti-predator behavior

For protection from land predators, the plains zebra retreats into open areas with good visibility at night. When the groups forage or sleep, one zebra will keep watch, and if a predator is spotted, it will bark or snort loudly.^[8] When being hunted by hyenas or wild dogs, a zebra harem stays close together and cooperates to protect threatened members,^[4] particularly the young. The harem stallion will go on the offensive and attack the dogs or hyenas.^[4] Though hyenas may harass the stallion, they usually only concentrate on the herd, and attempt to dodge the stallion's assaults. Unlike stallions, mares typically only react aggressively to hyenas or dogs when their foals are threatened. Unlike wildebeest, zebras rarely take to water when escaping hyenas.^[22] With lions, a zebra's best defense is to outpace them, as lions do not have as much endurance as hyenas or wild dogs. Cheetahs and leopards are mostly threats to foals as an adult zebra is fully capable of driving them away.

Human interactions

Conservation

Overall, the plains zebra population remains stable, and the species faces no major threat that would cause range-wide decline.^[1] The zebra can be found in numerous protected areas across its range, including the Serengeti National Park in Tanzania, Tsavo and Masai Mara in Kenya, Hwange National Park in Zimbabwe, Etosha National Park in Namibia, and Kruger National Park in South Africa. There are some stable populations in unprotected areas.^[1]

Zebras on the Botswana coat of arms

Some local populations, though, have faced great declines and even extinctions. One subspecies, the quagga, is now extinct. In Tanzania, the zebra population has decreased by 20% from the late 1990s to the mid 2000s.^[1] Zebras are threatened by hunting for their hide and meat, and habitat change from farming. They also compete with livestock for food,^[23][24] and are sometimes culled. Poaching is largely a threat to northern populations, while southern populations are threatened mostly by habitat loss. Recent civil wars in Rwanda, Somalia, Sudan, Ethiopia, and Uganda have caused dramatic declines in all wildlife populations, including those of plains zebra. It is now extinct in Burundi. Civil war in Angola during much of the past 25 years has devastated its wildlife populations, including its once-abundant plains zebra, and destroyed the national parks administration and infrastructure.

Nevertheless, plains zebras are protected in most of their range. They are an important economic source in tourism.

In culture

The zebra is revered in some African cultures as a symbol of beauty. In the dances of the Karamojong tribe of Uganda, women would paint themselves in zebra stripes and act like them.^[8] The Dube tribe of South Africa features a zebra on its totem. Zebras also appear on the coat of arms of Botswana.

References

Notes

1. ^ ^{a b c d e} Hack, M.A and Lorenzen, E. (2008). Equus quagga. In: IUCN 2008. IUCN Red List of Threatened Species. Downloaded on 10 April 2009. Database entry includes a brief justification of why this species is of least concern.
2. ^ Grubb, Peter (16 November 2005). "Order Perissodactyla (pp. 629-636)". In Wilson, Don E., and Reeder, DeeAnn M., eds. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press, 2 vols. (2142 pp.). p. 630. ISBN 978-0-8018-8221-0. OCLC 62265494. http://www.bucknell.edu/msw3/browse.asp?id=14100008. 
3. ^ Orlando, Ludovic; et al. (2009). "Revising the recent evolutionary history of equids using ancient DNA". PNAS 106: 21754–21759. 
4. ^ ^{a b c d e f g h i j k l m} Estes 1991
5. ^ J. E. Cordingley, S. R. Sundaresan, I. R. Fischhoff, B. Shapiro, J. Ruskey, D. I. Rubenstein (2009). Is the endangered Grevy's zebra threatened by hybridization?. Animal Conservation. 12: 505 - 513.
6. ^ [1] (2011).
7. ^ [2] (2011).
8. ^ ^{a b c d e f g h i} Kingdon 1979
9. ^ ^{a b c d e} Moehlman 2002
10. ^ Mutations
11. ^ Mount Kenya Bush Drums December 2006
12. ^ Groves C. P. (1974) Horses, Asses and Zebras in the Wild. Ralph Curtis Books, Hollywood
13. ^ Lamprey, H. F. (1963). Ecological separation of large mammal species in the Tangayika Game Reserve, Tangayika. E. Afr. Wildl. J. 63–93
14. ^ Moehlman 2003
15. ^ Wilderness Safaris - Zebra attack Blue Wildebeest foal
16. ^ National Geographic Zebra: Patterns in the Grass (1991)
17. ^ ^{a b c} Rubenstein and Hack
18. ^ ^{a b c d} Moss 1982
19. ^ ^{a b c} Klingel 1969
20. ^ ^{a b} Adlen et al. 1995
21. ^ Further evidence for male infanticide and feticide in captive plains zebras
22. ^ Kruuk, Hans (1972). The Spotted Hyena: A study of predation and social behaviour. p. 335. ISBN 0563208449. 
23. ^ Young, T.P.; T.M. Palmer & M.E. Gadd (2005). "Competition and compensation among cattle, zebras, and elephants in a semi-arid savanna in Laikipia, Kenya". Biological Conservation 121: 351–359. 
24. ^ Odadi, W.O.; T.P. Young & J.B. Okeyo-Owour (2009). "The effects of wild herbivores on cattle intake and movement rates in Laikipia rangeland, Kenya.". Applied Animal Behaviour Science 116: 120–125. 

General references

• Alden, P. C., Estes, R.D., Schlitter, D., McBride, B. (1995). National Audubon Society Field Guide to African Wildlife. New York, Chanticleer Press, Inc. pg. 151
• Estes, R. (1991). The Behavior Guide to African Mammals, Including Hoofed Mammals, Carnivores, Primates. Los Angeles, University of California Press. pgs. 242–246
• Groves, C.P. and Bell, H.B. 2004. New investigations on the taxonomy of the zebras genus Equus, subgenus Hippotigris. Mammalian Biology. 69: 182–196.
• "Equus quagga ssp. quagga". IUCN Red List of Threatened Species. Version 2011.1. International Union for Conservation of Nature. 2008. http://www.iucnredlist.org/apps/redlist/details/7957. Retrieved 17 August 2011.  (Extinct subspecies of the Plains Zebra.)
• Higuchi et al. (1987). Mitochondrial DNA of the Extinct Quagga: Relatedness and Extent of Postmortem Change. Journal of Molecular Evolution 25:283–287.
• Kingdon, J. (1979). East African Mammals: An Atlas of Evolution in Africa, Volume 3, Part B: Large Mammals. Chicago, University of Chicago Press. pgs. 165–179
• Klingel, H., (1969). Reproduction in the plains zebra Equus burchelli boehmi: behaviour and ecological factors. J. Reprod. Fertil., Suppl. 6: 339–345.
• Moelman, P.D. 2002. Equids. Zebras, Assess and Horses. Status Survey and Conservation Action Plan. IUCN/SSC Equid Specialist Group. IUCN, Gland, Switzerland. Chapter 4. Status and Action Plan for the Plains Zebra (Equus burchelli). Mace A. Hack, Rod East and Dan J Rubenstein. pgs. 43–57.
• Moehlman, P. D. (2003). Grizmek's Animal Life Encyclopedia. Mammals IV. Detroit, The Gale Group, Inc. 15.
• Moss, C., Ed. (1982). Portraits in the Wild, Animal Behavior in East Africa. Chicago, University of Chicago Press.
• Rubenstein, D. I. & M. Hack (2004) Natural and sexual selection and the evolution of multi-level societies: insights from zebras with comparisons to primates. pp. 266–279. In: Sexual Selection in Primates: New and Comparative Perspectives. P. Kappeler and C. P. van Schaik (eds.). Cambridge University Press.