Editing Plains zebra

{{Short description|Species of zebra}}
{{Good article}}
{{Speciesbox
| fossil_range = [[Pleistocene]] – Recent<ref>{{Cite web|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=90653|title = Fossilworks: Equus quagga}}</ref>
| image = Equus quagga burchellii - Etosha, 2014.jpg
| image_caption = [[Burchell's zebra]] (''E. quagga burchellii'') at [[Etosha National Park]] in [[Namibia]]
| status = NT
| status_system = IUCN3.1
| status_ref = <ref name="iucn status 19 November 2021">{{cite iucn |author=King, S.R.B. |author2=Moehlman, P.D. |date=2016 |title=''Equus quagga'' |volume=2016 |page=e.T41013A45172424 |doi=10.2305/IUCN.UK.2016-2.RLTS.T41013A45172424.en |access-date=19 November 2021}}</ref>
| genus = Equus
| parent = Equus (Hippotigris)
| species = quagga
| authority = [[Pieter Boddaert|Boddaert]], 1785
| synonyms_ref = <ref name="iucn status 19 November 2021" />
| synonyms = * ''Equus burchelli'' <small>(Gray, 1824) [orth. error]</small>
* ''Equus burchellii'' <small>Schinz, 1845</small>
| subdivision_ranks = Subspecies
| subdivision = ''[[Quagga|E. q. quagga]]'' <br />
''[[Burchell's zebra|E. q. burchellii]]''<br />
''[[Grant's zebra|E. q. boehmi]]''<br />
''[[Maneless zebra|E. q. borensis]]''<br />
''[[Chapman's zebra|E. q. chapmani]]''<br />
''[[Crawshay's zebra|E. q. crawshayi]]''<br />
''[[Selous' zebra|E. q. selousi]]''
| range_map = Plains Zebra area.png
| range_map_caption = {{legend0|#9f1105|&nbsp;plains zebra range}}
}}
The '''plains zebra''' ('''''Equus quagga''''', formerly ''Equus burchellii'') is the most common and geographically widespread species of [[zebra]]. Its range is fragmented, but spans much of southern and eastern Africa south of the [[Sahara]]. Six or seven [[subspecies]] have been recognised, including the [[quagga]] which was thought to be a separate species. More recent research supports variations in zebra populations being [[Cline (biology)|cline]]s rather than subspecies.

Plains zebras are intermediate in size between the larger [[Grévy's zebra]] and the smaller [[mountain zebra]] and tend to have broader stripes than both. Great variation in coat patterns exists between clines and individuals. The plains zebra's habitat is generally, but not exclusively, treeless grasslands and savanna woodlands, both tropical and temperate. They generally avoid desert, dense rainforest and permanent wetlands. Zebras are preyed upon by [[lion]]s and [[spotted hyena]]s, [[Nile crocodile]]s and, to a lesser extent, [[African leopard|leopards]], [[cheetah]]s and [[African wild dog]]s.

Plains zebras are a highly social species, forming harems with a single [[stallion]], several [[mare]]s and their recent offspring; bachelor groups also form. Groups may come together to form herds. The animals keep watch for predators; they bark or snort when they see a predator and the harem stallion attacks predators to defend his harem.

The plains zebra remains common in [[game reserve]]s, 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]]. The loss of open grasslands due to [[woody plant encroachment]] increases predation risk and therewith habitat.<ref name=":0">{{Cite journal |last1=Chen |first1=Anping |last2=Reperant |first2=Leslie |last3=Fischhoff |first3=Ilya R. |last4=Rubenstein |first4=Daniel I. |date=2021-07-01 |title=Increased vigilance of plains zebras (Equus quagga) in response to more bush coverage in a Kenyan savanna |journal=Climate Change Ecology |volume=1 |pages=100001 |doi=10.1016/j.ecochg.2021.100001 |issn=2666-9005|doi-access=free |bibcode=2021CCEco...100001C }}</ref> Plains zebra are listed as [[near threatened]] by the [[IUCN]] as of 2016. The species population is stable and not endangered, though populations in most countries have declined sharply.<ref>{{Cite report |url=http://www.iucnredlist.org/details/41013/0 |title=Equus quagga: King, S.R.B. & Moehlman, P.D.: The IUCN Red List of Threatened Species 2016: e.T41013A45172424 |last=IUCN |date=2016-06-20 |publisher=International Union for Conservation of Nature |doi=10.2305/iucn.uk.2016-2.rlts.t41013a45172424.en |language=en}}</ref>

==Taxonomy==
The plains zebra was formally classified by British zoologist [[John Edward Gray]] in 1824 as ''Equus burchellii''. After the quagga, described by [[Pieter Boddaert]] in 1785, was found to be the same species in the 21st century, the plains zebra was reclassified as ''Equus quagga'' due to the [[principle of priority]].<ref>{{cite journal|author=Heywood, P.|year=2015|title=The Micro-politics of Macromolecules in the Taxonomy and Restoration of Quaggas|journal=Kronos|volume=41|issue=1|pages=314–337|url=http://www.scielo.org.za/scielo.php?script=sci_arttext&pid=S0259-01902015000100013}}</ref> The plains zebra and [[mountain zebra]] were traditionally placed in the [[subgenus]] ''Hippotigris'', in contrast to [[Grévy's zebra]], which was considered the sole species of the subgenus ''Dolichohippus'';<ref name="Prothero 2003">{{Cite book|title=Horns, Tusks, and Flippers: The Evolution of Hoofed Mammals'|author1=Prothero D.R |author2=Schoch R. M |url=https://books.google.com/books?id=kWpQX-sfsLgC&q=Horns,+Tusks,+and+Flippers:+The+Evolution+of+Hoofed+Mammals|publisher=Johns Hopkins University Press|pages=216–18|year=2003|isbn=978-0-801-87135-1}}</ref> however, recent (2013) [[phylogenetic]] evidence finds that plains zebras are more closely related to Grévy's zebras than mountain zebras.<ref name=Vilstrup>{{cite journal|author=Vilstrup, Julia T.|year=2013|title=Mitochondrial Phylogenomics of Modern and Ancient Equids|journal=PLOS ONE|volume=8|issue=2|page=e55950|doi=10.1371/journal.pone.0055950|display-authors=etal|pmid=23437078|pmc=3577844|bibcode=2013PLoSO...855950V|doi-access=free}}</ref> Groves and Bell (2004) place all three species in the subgenus ''Hippotigris'',<ref name="GrovesBell2004">{{cite journal|author=Groves, C. P.; Bell, C. H.|year=2004|title=New investigations on the taxonomy of the zebras genus ''Equus'', subgenus ''Hippotigris''|journal=Mammalian Biology|volume=69|issue=3|pages=182–196|doi=10.1078/1616-5047-00133|bibcode=2004MamBi..69..182G }}</ref> and zebras appear to be a [[monophyletic]] lineage.<ref name=Vilstrup/> In areas where plains zebras are [[sympatric]] with Grévy's zebras, finding them in the same herds is not unusual,<ref name="Estes 1991">{{cite book|author=Estes, R.|year=1991|title=The Behavior Guide to African Mammals, Including Hoofed Mammals, Carnivores, Primates|url=https://archive.org/details/isbn_0520080858|url-access=registration|publisher=University of California Press|pages=[https://archive.org/details/isbn_0520080858/page/242 242–246]|isbn=978-0-520-08085-0}}</ref> and [[Zebroid|fertile hybrids]] occur.<ref>{{cite journal|author=J. E. Cordingley, S. R. Sundaresan, I. R. Fischhoff, B. Shapiro, J. Ruskey, D. I. Rubenstein|year=2009|title=Is the endangered Grevy's zebra threatened by hybridization?|journal=Animal Conservation|volume=12|issue=6|pages=505–513|doi=10.1111/j.1469-1795.2009.00294.x|bibcode=2009AnCon..12..505C |s2cid=18388598}}</ref>

===Subspecies===
In their 2004 study of cranial and pelage differences between specimens, Groves and Bell found support for the division of the plains zebra into six subspecies:<ref name="GrovesBell2004"/>
{| class="wikitable "
|-
! Image !! Subspecies !!  Distribution
|-
|[[File:Maneless zebras at Kidepo Valley NP - Uganda.jpg|150px]]||[[Maneless zebra]], ''Equus quagga borensis'' – Lönnberg, 1921 ||  Northwestern [[Kenya]] (from [[Uasin Gishu County|Uasin Gishu]] and [[Lake Baringo]]) to the [[Karamoja]] district of [[Uganda]].
|-
|[[File:Equus quagga.jpg|150px]]||[[Grant's zebra]], ''Equus quagga boehmi'' – Matschie, 1892 ||  [[Zambia]] west of the [[Luangwa River]] and west to Kariba, [[Katanga Province]] of the [[Democratic Republic of the Congo]], north to the Kibanzao Plateau, and in [[Tanzania]] north from Nyangaui and [[Kibwezi]] into southwestern Kenya as far as [[Sotik Constituency|Sotik]] and southwestern [[Ethiopia]] from [[Omo National Park|Omo Valley]] to [[Nechisar National Park|Nechisar plains]] in the north and [[Yabelo Wildlife Sanctuary|Yabello]] and [[Borena National Park|Borena Zone]] to the east. 
|-
|[[File:Zebra Side.JPG|150px]]||[[Crawshay's zebra]], ''Equus quagga crawshayi'' – De Winton, 1896 || Eastern Zambia, east of the Luangwa River, [[Malawi]], southeastern Tanzania, and northern [[Mozambique]] south to the Gorongoza District
|-
|[[File:Burchell's Zebra (Equus quagga burchellii) (15844593253).jpg|150px]]|| [[Chapman's zebra]], ''Equus quagga chapmani'' – Layard, 1865 ||  North-east [[South Africa]], north to Zimbabwe, west into [[Botswana]], the [[Caprivi Strip]] in [[Namibia]], and southern [[Angola]].
|-
|[[File:Equus quagga burchellii - Etosha, 2014.jpg|150px]]<!--Common zebra 1.jpg-->|| [[Burchell's zebra]], ''Equus quagga burchellii'' – Gray, 1824 || Southern Botswana to Etosha and the [[Kaokoveld]], and southeast to [[Eswatini]] and [[KwaZulu-Natal]]. 
|-
|[[File:Quagga Naturhistorisches Museum Basel 27102013 3.jpg|150px]]|| [[Quagga]], [[Extinction|†]]''Equus quagga quagga'' – Boddaert, 1785||  South Africa 
|-
|[[File:Zeitschrift für wissenschaftliche Zoologie (1906) (14762972812).jpg|150px]]|| [[Selous' zebra]], ''Equus quagga selousi'' – Pocock, 1897*|| Mozambique
|-
|}
<nowiki>*</nowiki>Sometimes a seventh subspecies is recognised.<ref>{{cite book|author=Beolens, B.; Watkins, M.; Grayson, M.|year=2009|title=The Eponym Dictionary of Mammals|publisher=Johns Hopkins University Press|pages=167|isbn=978-0801893049}}</ref>

Burchell's zebra was thought to have been hunted to extinction. However, Groves and Bell concluded that "the extinct true Burchell's zebra is a phantom". Careful study of the original zebra populations in Zululand and Eswatini 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'' 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 suggested that the correct name for the subspecies must be ''burchellii'', not ''antiquorum''.<ref name="GrovesBell2004"/>

A 2005 genetic study confirmed the quagga being the same species as the plains zebra. It showed that the quagga had little genetic diversity and that it diverged from the other plains zebra subspecies only 120,000–290,000 years ago, during the [[Pleistocene]] and, possibly, the penultimate glacial maximum. Its distinct coat pattern may have evolved rapidly because of geographical isolation and/or adaptation to a drier environment. In addition, plains zebra subspecies tend to have less striping the further south they live, and the quagga was the most southern-living of them all.<ref name="smithsonian">{{Cite journal |last1=Hofreiter |first1=M. |last2=Caccone |first2=A. |last3=Fleischer |first3=R. C. |last4=Glaberman |first4=S. |last5=Rohland |first5=N. |last6=Leonard |first6=J. A. |doi=10.1098/rsbl.2005.0323 |title=A rapid loss of stripes: The evolutionary history of the extinct quagga |journal=Biology Letters |volume=1 |issue=3 |pages=291–295 |year=2005 |pmid=17148190 |pmc=1617154}}</ref>

The simplified [[cladogram]] below is based on the 2005 analysis (some taxa shared [[haplotype]]s and could therefore not be differentiated):<ref name="smithsonian"/>
{{clade| style=font-size:100%; line-height:100%
|1={{clade
 |1=[[Mountain zebra]] (''E. zebra'')
 |2={{clade
  |1=[[Grévy's zebra]] (''E. grevyi'')
  |2={{clade
    |1={{clade
     |1='''Quagga''' ('''''E. q. quagga''''')
     |2=[[Damara zebra]] (''E. q. antiquorum'')-[[Chapman's zebra]] (''E. q. chapmani'')}}
    |2=[[Grant's zebra]] (''E. q. boehmi'') }}
  }}
 }}
}}

A 2018 DNA study found no evidence for a subspecies structure in plains zebras but, instead, observed a north–south genetic continuum. Modern plains zebra populations appear to have originated from Southern Africa around 370,000 years ago with plains zebras in Uganda, the most northern population, being the most distinct.<ref name="PedersenAlbrechtsen2018">{{cite journal |last1=Pedersen |first1=Casper-Emil T. |last2=Albrechtsen |first2=Anders |last3=Etter |first3=Paul D. |last4=Johnson |first4=Eric A. |last5=Orlando |first5=Ludovic |last6=Chikhi |first6=Lounes |last7=Siegismund |first7=Hans R. |last8=Heller |first8=Rasmus |year=2018|title=A southern African origin and cryptic structure in the highly mobile plains zebra |journal=Nature Ecology & Evolution |volume=2 |issue=3 |pages=491–498 |issn=2397-334X |doi=10.1038/s41559-017-0453-7 |pmid=29358610 |bibcode=2018NatEE...2..491P |s2cid=3333849 }}</ref>

==Physical description==
[[File:Equus burchelli - Kunming Natural History Museum of Zoology - DSC02371.JPG|thumb|left|222px|The skeleton exhibit in the [[Kunming Natural History Museum of Zoology]], [[Kunming]], [[Yunnan]], China]]
The plains zebra stands at a height of {{cvt|127|–|140|cm}} with a head-body length of {{cvt|217|–|246|cm}} and a tail length of {{cvt|47|–|56.5|cm}}. Males weigh {{cvt|220|–|322|kg}} while females weigh {{cvt|175|–|250|kg}}.<ref name="Kingdon 1988">{{cite book|author=Kingdon, J.|year=1988|title=East African Mammals: An Atlas of Evolution in Africa, Volume 3, Part B: Large Mammals|publisher=University of Chicago Press|pages=165–179|isbn=978-0-226-43722-4}}</ref> The species is intermediate in size between the larger Grévy's zebra and the smaller mountain zebra. It is dumpy bodied with relatively short legs and a skull with a convex forehead and a somewhat concave nose profile. The neck is thicker in males than in females. The ears are upright and have rounded tips. They are shorter than in the mountain zebra and narrower than in Grévy's zebra. As with all wild equids, the plains zebra has an erect mane along the neck and a tuft of hair at the end of the tail.<ref name="Grub 1981">{{cite journal | last1 = Grubb | first1 = P. | year = 1981 | title = Equus burchellii | journal = Mammalian Species | issue = 157| pages = 1–9 | doi=10.2307/3503962| jstor = 3503962 | doi-access = free }}</ref><ref>{{Cite book|author=Rubenstein, D. I.|contribution=Horse, Zebras and Asses|year=2001|title=The Encyclopedia of Mammals |edition= 2nd|editor=MacDonald, D. W.|publisher=Oxford University Press|page=468|isbn=978-0-7607-1969-5}}</ref><ref name="Skinner"/> The body hair of a zebra is {{cvt|9.4|±|4|mm}},<ref name="Skinner"/> shorter than in other African ungulates.<ref name="Caro">{{cite journal|author1=Caro, T.|author2=Izzo, A.|author3=Reiner, R. C.|author4=Walker, H.|author5=Stankowich, T |year=2014 |title= The function of zebra stripes|journal=Nature Communications |volume=5 |pages=3535 |doi=10.1038/ncomms4535 |pmid=24691390 |bibcode=2014NatCo...5.3535C |author1-link=Tim Caro|doi-access=free}}</ref>

[[File:Zebras in Tanzania 4310 Nevit.jpg|thumb|right|Close-up of striping pattern of zebra in Tanzania]]
Like all zebras, they are [[zebra stripes|boldly striped in black and white]] and no two individuals look exactly alike. Compared to other species, the plains zebra has broader stripes. The stripes are vertical on the fore part of the body, and tend towards the horizontal on the hindquarters. Northern zebra populations have narrower and more defined striping; southern populations have varied but lesser amounts of striping on the under parts, the legs and the hindquarters. Southern populations also have brown "shadow" stripes between the black and white colouring. These are absent or poorly expressed in northern zebras. The natal coat of a [[foal]] is brown and white and the brown darkens with age.<ref name="Grub 1981"/><ref name="Kingdon 1988"/><ref name="Moehlman 2002">{{Cite book |last1=Hack |first1=Mace A. |url=https://archive.org/details/equidszebrasasse0000unse |title=Equids. Zebras, Asses and Horses. Status Survey and Conservation Action Plan |last2=East |first2=Rod |last3=Rubenstein |first3=Dan J. |publisher=IUCN/SSC Equid Specialist Group. IUCN |year=2002 |isbn=9782831706474 |editor-last=Moehlman |editor-first=P. D. |pages=43–57 |chapter=Status and Action Plan for the Plains Zebra (''Equus burchelli'') |chapter-url=https://archive.org/details/equidszebrasasse0000unse/page/43 |url-access=limited}}</ref>

Various abnormalities of the patterns have been documented in plains zebras. [[Melanistic]] zebras have high concentrations of dark stripes on the torso but low concentrations on the legs. "Spotted" individuals display interruptions in black striping patterns.<ref name=Larison2020>{{cite journal|last1=Larison|first1=Brenda|last2=Kaelin|first2=Christopher B.|last3=Harrigan|first3=Ryan|display-authors=etal|year=2020|title=Population structure, inbreeding and stripe pattern abnormalities in plains zebras|journal=Molecular Ecology|volume=30|issue=2|pages=379–390|doi=10.1111/mec.15728|pmid=33174253|s2cid=226305574}}</ref> There have even been [[Polymorphism (biology)|morphs]] with white spots on dark backgrounds.<ref>{{Cite book |title=Zebra Stripes |last=Caro |first=Tim |publisher=[[University of Chicago Press]]|page=20|year=2016 |isbn=978-0-226-41101-9 |url=https://books.google.com/books?id=3o-EDQAAQBAJ&q=zebra+stripes}}</ref> Striping abnormalities have been linked to [[inbreeding]].<ref name=Larison2020/> [[Albino]] zebras have been recorded in the forests of [[Mount Kenya]], with the dark stripes being blonde.<ref>{{cite web |url=https://www.nationalgeographic.com/animals/2019/03/rare-partially-albino-zebra-spotted-in-serengeti/ |archive-url=https://web.archive.org/web/20190329161519/https://www.nationalgeographic.com/animals/2019/03/rare-partially-albino-zebra-spotted-in-serengeti/ |url-status=dead |archive-date=March 29, 2019 |title=Extremely Rare 'Blonde' Zebra Photographed |publisher=[[National Geographic]] |date=29 March 2019 |access-date=25 May 2020}}</ref> The quagga had brown and white stripes on the head and neck, brown upper parts and a white belly, tail and legs.<ref name="Walker">{{cite book |last=Nowak |first=R. M. |year=1999 |title=Walker's Mammals of the World |volume=1 |publisher=[[Johns Hopkins University Press]] |pages=1024–1025 |isbn=978-0-8018-5789-8}}</ref>

==Ecology==

===Range and habitat===
[[File:Maasai Mara 341.jpg|thumb|right|Zebras migrating with [[wildebeest]] in the [[Maasai Mara]]]]
The plains zebra's range stops short of the Sahara from [[South Sudan]] and southern [[Ethiopia]] extending south along eastern Africa, as far as Zambia, Mozambique, and Malawi, before spreading into most southern African countries. They may have lived in [[Algeria]] in the [[Neolithic era]]. Plains zebras generally live in treeless grasslands and savanna woodlands, 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.<ref name="Moehlman 2002"/> They generally prefer [[Acacieae]] woodlands over ''[[Commiphora]]''. They are water-dependent and live in more [[Mesic habitat|mesic]] environments than other African equids. They seldom wander {{convert|10|–|12|km|abbr=on}} from a water source. Zebras also live in elevations from [[sea level]] to {{convert|4,300|ft|m}} on [[Mount Kenya]].<ref name="Estes 1991"/><ref name="Grub 1981"/><ref name="Skinner"/>

Depending on the population, zebra herds may be sedentary, being highly dense with small ranges, or [[Animal migration|migratory]], being less populated with separate, extensive dry and wet home ranges.<ref name="Grub 1981"/> When migrating, zebras appear to rely on some memory of the locations where foraging conditions were best and may predict conditions months before their arrival.<ref>{{cite journal |author=Bracis, C.; Mueller, T.|year=2017|title=Memory, not just perception, plays an important role in terrestrial mammalian migration |journal=Proceedings of the Royal Society B: Biological Sciences |volume=284 |issue=1855 |page=20170449 |doi=10.1098/rspb.2017.0449 |pmid=28539516|pmc=5454266}}</ref>

The loss of open grasslands due to [[woody plant encroachment]] increases predation risk and therewith habitat.<ref name=":0" />

===Diet and predation===
Plains zebras primarily feed on [[grass]]; preferred species being ''[[Themeda triandra]]'', ''[[Cynodon dactylon]]'', ''[[Eragrostis superba]]'' and ''[[Cenchrus ciliaris]]''. Zebra sometimes [[Browsing (herbivory)|browse]] or dig for [[corm]]s and [[rhizome]]s during the dry season.<ref name="Grub 1981"/> They appear to partial to eating scorched ''[[Colophospermum mopane]]'' and ''[[Pterocarpus rotundifolius]]'', consuming both the leaves and twigs.<ref name="Skinner"/>

Plains zebras are adapted for grazing on both long, tough grass stems and newly emerging short grass. In some areas, it rarely feeds below {{convert|100|–|150|mm|abbr=on}} to ground level. It ranges more widely than many other species, even into [[woodland]]s, and it is often the first grazing species to appear in a well-vegetated area.<ref name="Estes 1991"/><ref name="Skinner"/> The flexible upper lip allows them to push plant material between the incisors to cut.<ref name="Skinner"/> Zebras have a less efficient digestive system than [[ruminant]]s but food passage is twice as fast.<ref name="Grub 1981"/> 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 specialised grazers such as [[blue wildebeest]]s and [[Thomson's gazelle]]s.<ref name="Estes 1991"/>

[[File:Lions and a Zebra b.jpg|thumb|Lions feeding on a zebra]]
The plains zebra's major predators are [[lion]]s and [[spotted hyena]]s. Lions are most successful when targeting lone individuals, usually an old male while hyenas chase and isolate an individual from the group, usually a female or foal.<ref name="Kingdon 1988"/> [[Nile crocodile]]s also prey on zebras when they are near water.<ref>{{cite book|author=Kennedy, A. S., & Kennedy, V.|year=2013|title=Animals of the Masai Mara|publisher=Princeton University Press|page=130|isbn=978-0691156019}}</ref> Less common predators include [[leopard]]s, [[cheetah]]s and [[African wild dog]]s, which mostly hunt foals.<ref name="Grub 1981"/> When in the presence of a lion, zebras remain alert and stand in a semi-circle at as much as {{convert|100|m|ft|abbr=on}} and no less than {{convert|50|m|ft|abbr=on}}. Stallions sometimes try to drive lions away with bluff charges. By contrast, zebras may approach cheetahs and wild dogs and a single hyena is allowed to come within a few metres. To escape from predators, an adult zebra can run at {{convert|60|-|70|km/h|abbr=on}}.<ref name="Kingdon 1988"/> When being hunted by hyenas or wild dogs, a zebra harem stays close together and cooperates to protect threatened members, particularly the young. The harem stallion goes on the offensive and attacks the dogs or hyenas.<ref name="Estes 1991"/>

==Behaviour==
Plains zebras are nomadic and non-territorial, [[home range]]s vary from {{convert|30|km2|abbr=on}} to {{convert|600|km2|abbr=on}}, depending on the area and if the population is migratory. They are more [[Diurnality|active during the day]] and spend most of their time feeding. Other activities include [[dust bathing]], rubbing, drinking and intermittent resting which is very brief. At night, zebra activity is subdued except when threatened by predators. They may rest or sleep laying down, while one individual keeps guard.<ref name="Estes 1991"/>

===Social structure===
[[File:Zebra Botswana edit02.jpg|thumb|A zebra harem within a herd]]
The plains zebra is highly social and usually forms small family groups called [[Harem (zoology)|harem]]s, 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 "[[Bachelor herd|bachelors]]" also exist. These are stable groups of up to 15 males with an age-based hierarchy, led by a young male. These males stay in their groups until they are ready to start a harem. The bachelors prepare for their adult roles with play fights and greeting/challenge rituals, which take up most of their activities.<ref name="Estes 1991"/>

Multiple harems and bachelor groups come together to form larger herds of hundreds of animals, especially during migrations. Plains zebras are unusual among harem-holding species in forming these groups. In addition, pairs of harems may create temporarily stable subgroups within a herd, allowing individuals to interact with those outside their group. Among harem-holding species, this has only been observed in primates such as the [[gelada]] and the [[hamadryas baboon]].<ref name="Rubenstein and Hack 2004">{{cite book|author=Rubenstein, D. I.; Hack, M.|year=2004|contribution=Natural and sexual selection and the evolution of multi-level societies: insights from zebras with comparisons to primates|pages=266–279|title=Sexual Selection in Primates: New and Comparative Perspectives|editor=Kappeler, P.; van Schaik, C. P.|publisher=Cambridge University Press|isbn=978-0521537384}}</ref> Bachelor groups tend to be at the periphery of herds and when the herd moves, the bachelors trail behind.<ref name="Skinner"/>

Stallions form and expand their harems by abducting young mares from their natal harems.<ref name="Estes 1991"/> When a mare reaches sexual maturity and has her first [[oestrous]] cycle, she attracts the attention of nearby stallions, both bachelors and harem leaders. Her family stallion (likely her father) chases off or fights stallions attempting to abduct her. Even after a young mare is isolated from her natal harem, the fight over her continues until her oestrous cycle is over and it starts again with the next oestrous cycle. It is rare that the mare's original abductor keeps her for long.<ref name="Klingel 1969">{{cite journal | last1 = Klingel | first1 = H. | year = 1969 | title = Reproduction in the plains zebra ''Equus burchelli boehmi'': behaviour and ecological factors | journal = J. Reprod. Fertil. Suppl. | volume = 6 | pages = 339–345 }}</ref> When the mare finally [[Ovulation|ovulates]], the male that impregnates her keeps her for good. Thus, the mare becomes a permanent member of a new harem.<ref name="Klingel 1969"/><ref name="Adlen et al. 1995">{{cite book|author=Alden, P. C., Estes, R. D., Schlitter, D., McBride, B.|year=1995|title=National Audubon Society Field Guide to African Wildlife|publisher=Chanticleer Press, Inc|page=151|isbn=978-0679432340}}</ref> Oestrus in a female becomes less noticeable to outside males as she gets older, hence competition for older females is virtually nonexistent.<ref name="Grub 1981"/>

[[File:Fighting Burchell's Zebras in Addo National Elephant Park.JPG|thumb|left|Two male zebras fighting in the [[Addo Elephant National Park]], South Africa]]
Mares exist in a hierarchy, with the alpha female being the first to mate with the harem 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 harem stallion must shield the new mares until the aggression subsides.<ref name="Estes 1991"/><ref name="Adlen et al. 1995"/> The most recently added females rank lowest. Females that become unfit or weak may drop in their rank, though. The female membership of a harem stays intact, even if a new stallion takes over. During herd gatherings, family stallions may be cordial towards each other, while the mares are less tolerant.<ref name="Grub 1981"/>

A stallion defends his harem from other males. When challenged, the stallion issues 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 lies still on the ground as if surrendering, but once the other male lets up, he strikes and continues the fight. Most fighting occurs over young mares in oestrus and as long as a harem stallion is healthy, he usually is not 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.<ref name="Estes 1991"/>

===Communication===
[[File:Equus quagga (185908025).jpg|thumb|right|Zebras mutually grooming]]
At least six different calls have been documented for the plains zebra. One is its distinctive, high-pitched, contact call (commonly called "[[bark (sound)|barking]]") heard as "a-''ha'', a-''ha'', a-''ha''" or "kwa-ha, kaw-ha, ha, ha"<ref name="Grub 1981"/> also transcribed as "kwahaah",<ref name=Max>{{cite news |last=Max|first=D. T. |title=Can You Revive an Extinct Animal? |url=https://www.nytimes.com/2006/01/01/magazine/01taxidermy.html |access-date=2014-03-03 |newspaper=[[The New York Times]] |date=1 January 2006}}</ref> or "oug-ga".<ref name="1911 quagga">{{EB1911 |noprescript=1 |wstitle=Quagga}}</ref> The species name ''quagga'' is derived from the [[Khoikhoi]] word for "zebra" and is [[Onomatopoeia|onomatopoeic]] for its call.<ref name=skinner>{{cite book |last1=Skinner |first1=J. D. |title=The Mammals of the Southern African Subregion |year=2005 |publisher=[[Cambridge University Press]] |isbn=978-0-521-84418-5 |pages=537–546 |edition = 3rd |last2=Chimimba |first2=C. T. |chapter=Equidae}}</ref> When a predator is sighted, a zebra makes a two-syllable alarm call. A loud snort is made when moving in cover of potential danger. When in contentment, a zebra makes a more drawn-out snort. Males make a short, high-pitched squeal when hurt, and foals  emit a drawn-out wail when in distress.<ref name="Grub 1981"/>

Two main facial expressions are made by zebras; the greeting and threat. In both cases, the lips are pulled back and chewing motions are made. Greeting involves the ears sticking up and directing forward; while the threat involves the ears down.<ref name="Grub 1981"/> Zebras strengthen their social bonds with [[Social grooming|grooming]]. Members of a harem nip and scrape along the neck, withers, and back with their teeth and lips. Mothers and foals groom the most often, followed by siblings. Grooming shows social status and eases aggressive behaviour.<ref name="Estes 1991"/>

===Reproduction and parenting===
[[File:Burchell's zebra (Equus quagga burchellii) suckling.jpg|thumb|Mother zebra nursing her foal]]
[[File:Grévy's × Plains Zebra, ol pejeta imported from iNaturalist photo 401473965 (cropped).jpg|thumb|Grévy's × plains zebra hybrid, alongside plains zebras.]]
The stallion mates with all his mares. Males exhibit the [[flehmen response]] to test for female receptivity, which involves the upper lip curling back to smell for urine (via the [[vomeronasal organ]]). The female signals her readiness for copulation by straddling her legs and raising her tail. The gestation period lasts around a year, and a single young is produced. Mares may give birth to one foal every twelve months. The birthing peak is during the rainy season. A mare gives birth within the vicinity of her group and while lying down on her side. The newborn foal weighs {{convert|30|–|35|kg|lb|abbr=on}} and the [[afterbirth]] is rarely consumed.<ref name="Skinner"/><ref name="Kingdon 1988"/><ref name="Grub 1981"/>

A newborn is capable of standing almost immediately and starts to eat grass within a week. Early on, a mother zebra keeps any other zebra away from her foal, including the stallion, the other mares, and even her previous offspring. Later, though, they all bond. Within the group, a foal has the same rank as its mother.<ref name="Grub 1981"/><ref name="Kingdon 1988"/><ref name="Nunez2011">{{cite book|author=Nunez, C. M. V.; Rubenstein, D. I.; Asa, C. S.|year=2011|contribution=Zebra Reproduction|title=Equine Reproduction|editor=McKinnon, A. O.; Squires, E. L.; Vaala, E. L.; Varner, D. D.|publisher=Wiley-Blackwell|pages=2851–2865|isbn=978-0813819716}}</ref><ref name="Skinner"/> The stallion is generally intolerant of foals that are not his, and zebras may practice [[Infanticide (zoology)|infanticide]] and [[feticide]].<ref>{{cite web |url=http://www.ivb.cz/folia/54/3/258-262.pdf |title=Further evidence for male infanticide and feticide in captive plains zebras |access-date=2012-07-03 |archive-url=https://web.archive.org/web/20120222174717/http://www.ivb.cz/folia/54/3/258-262.pdf |archive-date=2012-02-22 |url-status=dead }}</ref>

Mortality for foals is high in their first year of life and is usually caused by predation. However, zebra young are afforded more protection than those of species like wildebeest and [[hartebeest]]. A foal is usually weaned at around eleven months, but may suckle for longer. Females reach puberty before three years, and males after five or six. Young male zebras eventually leave their family groups as the relationship with their mothers fades after the birth of a sibling. The young stallion then seeks out other young stallions for company. Young females may stay in the harem until they are abducted by another stallion.<ref name="Kingdon 1988"/><ref name="Estes 1991"/><ref name="Nunez2011"/><ref name="Skinner">{{cite book|author=Skinner, J. D.; Chimimba, C. T.|year=2006|title=The Mammals of the Southern African Sub-region|publisher=Cambridge University Press|pages=243–246|isbn=978-0521844185}}</ref> Plains zebras have an average lifespan of 25 years in the wild.<ref>{{Cite web |date=2010-09-10 |title=Plains Zebra {{!}} National Geographic |url= https://www.nationalgeographic.com/animals/mammals/facts/plains-zebra |archive-url= https://web.archive.org/web/20210225171557/https://www.nationalgeographic.com/animals/mammals/facts/plains-zebra |url-status=dead |archive-date=February 25, 2021 |access-date=2022-12-15 |website=Animals |language=en}}</ref>

==Human interactions==
===Conservation===
[[Image:Quagga in enclosure.jpg|thumb|left|A quagga mare in London Zoo, 1870]]
In 2016, the plains zebra was classified as [[near threatened]] by the [[IUCN]]. As of that year, the total population is estimated to be around 500,000 individuals. The species remains common throughout its range but has experienced population declines in 10 of the 17 countries where it is native. They are stable in [[Ethiopia]], [[Malawi]], and [[South Africa]] and possibly [[Angola]]; stable or increasing in [[Mozambique]], [[Namibia]] and [[Eswatini]]; and decreasing in [[Botswana]], [[DR Congo]], [[Kenya]], [[Rwanda]], [[Somalia]], [[South Sudan]], [[Tanzania]], [[Uganda]], [[Zambia]], and [[Zimbabwe]]. They are extinct in [[Burundi]], [[Lesotho]] and possibly Somalia.<ref name="iucn status 19 November 2021" />

Zebras are threatened by hunting for their hide and meat, and habitat change from farming. They also compete with livestock for food,<ref name=comp>{{Cite journal | last = Young | first = T. P. |author2=T. M. Palmer |author3=M. E. Gadd | title = Competition and compensation among cattle, zebras, and elephants in a semi-arid savanna in Laikipia, Kenya | journal = Biological Conservation | volume = 121 | issue = 2 | pages = 351–359 | year = 2005 | doi=10.1016/j.biocon.2004.08.007| bibcode = 2005BCons.122..351Y }}</ref><ref name=foraging>{{Cite journal | last = Odadi | first = W. O. |author2=T. P. Young |author3=J. B. Okeyo-Owour | title = The effects of wild herbivores on cattle intake and movement rates in Laikipia rangeland, Kenya. | journal = Applied Animal Behaviour Science | volume = 116 | issue = 2–4 | pages = 120–125 | year = 2009 | doi=10.1016/j.applanim.2008.08.010}}</ref><ref name="Moehlman 2002"/> and fencing blocks migration routes.<ref name="iucn status 19 November 2021" /> Civil wars in some countries have also caused declines in zebra populations.<ref name="Moehlman 2002"/> 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. Some stable populations live in unprotected areas.<ref name="iucn status 19 November 2021" />

The quagga was hunted by early Dutch settlers and later by [[Afrikaner]]s to provide meat or for their skins. The skins were traded or used locally. The quagga was probably vulnerable to extinction due to its limited distribution, and it may have competed with domestic livestock for forage. The last known wild quagga died in 1878.<ref name="Weddell">{{cite book|author=Weddell, B. J. |year=2002 |title=Conserving Living Natural Resources: In the Context of a Changing World |url= https://archive.org/details/conservingliving00wedd_343|url-access=limited|publisher=Cambridge University Press|page=[https://archive.org/details/conservingliving00wedd_343/page/n62 46]|isbn=978-0-521-78812-0}}</ref> The last captive quagga, a female in Amsterdam's [[Natura Artis Magistra]] zoo, lived there from 9 May 1867 until it died on 12 August 1883, but its origin and cause of death are unclear.<ref name="ungulates">{{Cite journal |last=Van Bruggen |first=A.C. |year=1959 |title=Illustrated notes on some extinct South African ungulates |url=https://journals.co.za/doi/pdf/10.10520/AJA00382353_1382 |journal=South African Journal of Science |volume=55 |pages=197–200 }}</ref> In 1984, the quagga was the first extinct animal to have its DNA analysed, and the [[Quagga Project]] is trying to recreate the [[phenotype]] of hair coat pattern and related characteristics by selectively breeding Burchell's zebras.<ref name="Project">{{Cite journal |last1=Harley |first1=E.H. |last2=Knight |first2=M.H. |last3=Lardner |first3=C. |last4=Wooding |first4=B. |last5=Gregor |first5=M. |title=The Quagga Project: Progress over 20&nbsp;years of selective breeding |doi=10.3957/056.039.0206 |journal=South African Journal of Wildlife Research |volume=39 |issue=2 |pages=155 |year=2009 |citeseerx=10.1.1.653.4113 |s2cid=31506168 |url=http://quaggaproject.org/downloads/SAJWRpaper.pdf}}</ref>

===In popular culture===
[[File:Coat of arms of Botswana.svg|thumb|right|upright|Zebras on the [[Coat of arms of Botswana|Botswana coat of arms]]]]
Zebras have been featured in [[African art]] and [[African culture|culture]] for millennia. They have been depicted in [[rock art]] in Southern Africa (modern Botswana, Namibia and South Africa) dating from 20,000–28,000 years ago, though not as commonly as antelope species like [[Common eland|eland]]. How the zebra got its stripes has been the subject of [[Traditional African religions|folk tales]], some of which involve it being scorched by fire. The [[San people]] associated zebra stripes with water, rain and lighting due to its dazzling pattern.<ref name="reaktion"/>

[[File:Flag of Botswana.svg|thumb|left|upright|Black and white stripes on the [[Flag of Botswana|Botswana flag]] represent the stripes of a zebra.]]
The plains zebra is the [[national animal]] of the [[Botswana|Republic of Botswana]] and its stripes are depicted on the country's flag. The flag stripes also represent racial harmony in the country.<ref>{{cite web | title = Introducing the flag of Botswana | url = https://www.lonelyplanet.com/articles/flag-of-botswana-facts-history-and-design | date = 19 August 2020 | publisher = [[Lonely Planet]] | access-date = 9 August 2021 | archive-date = 9 August 2021 | archive-url = https://web.archive.org/web/20210809220239/https://www.lonelyplanet.com/articles/flag-of-botswana-facts-history-and-design | url-status = dead }}</ref> The zebra has also been associated with beauty and the women of various societies would paint much of their bodies in stripes. For the [[Shona people]] of Zimbabwe, the zebra is a [[totem]] animal, along with the eland, buffalo, lion and monkey. The zebra is praised in a poem as an "iridescent and glittering creature". Its stripes have symbolised the joining of male and female and at [[Great Zimbabwe]], zebra stripes decorate what is believed to be a ''domba'', a premarital school meant to initiate women into adulthood. In the [[Shona language]], the name "madhuve" means "woman/women of the zebra totem" and is a given name for girls in Zimbabwe.<ref name="reaktion">{{cite book|author1=Plumb, C. |author2=Shaw, S. |year=2018|title=Zebra|publisher=Reaktion Books|pages=37–46, 192–201|isbn=978-1780239354}}</ref>

Zebras have also been represented in Western culture. They have been thought of as a more exotic alternative to horse; the comic book character [[Sheena, Queen of the Jungle]] is depicted riding a zebra. The film ''[[Racing Stripes]]'' features a captive zebra ostracised from the horses and end up being ridden by a rebellious girl.<ref name="reaktion"/> In the film ''[[Fantasia (1940 film)|Fantasia]]'', two [[centaur]]s are depicted being half human and half zebra, instead of the typical half human and half horse.<ref>{{cite web |url= http://www.filmsite.org/fant.html|title= ''Fantasia'' (1940)|first= Tim|last= Dirks|date= n.d.|publisher= [[Filmsite.org]]|archive-url= https://web.archive.org/web/20140823181529/http://www.filmsite.org/fant.html|archive-date= 2014-08-23|url-status= live}}</ref> Zebras have been featured as characters in other animated films like ''[[Khumba]]'', ''[[The Lion King]]'' and the [[Madagascar (franchise)|''Madagascar'' films]].<ref name="reaktion"/>

==References==
{{Reflist|20em}}

==External links==
{{Wikispecies|Equus quagga}}
{{Commons category|Equus quagga}}

{{Equus|state=collapsed}}
{{Perissodactyla|state=collapsed}}
{{Taxonbar|from=Q245305}}
{{Authority control}}

[[Category:Zebras]]
[[Category:Fauna of East Africa]]
[[Category:Mammals of Southern Africa]]
[[Category:National symbols of Botswana]]
[[Category:Mammals described in 1785|plain zebra]]
[[Category:Taxa named by Pieter Boddaert]]