View source for Zebra

{{short description|Black-and-white striped animals in the equid family}}
{{distinguish|zebro}}
{{other uses}}
{{Featured article}}
{{pp-semi|small=yes}}
{{use British English|date=July 2020}}
{{use dmy dates|date=July 2020}}
{{Automatic taxobox
| fossil_range = {{Fossil range|2|0|earliest=2.3|ref=<ref name="Badenhorst2019"/>|PS=[[Pleistocene]]–[[Holocene|present]]}}
| image = Plains Zebra Equus quagga cropped.jpg
| image_alt = A herd of plains zebra ("Equus quagga")
| image_caption = A [[plains zebra]] (''Equus quagga'') among a herd in the [[Ngorongoro Crater]] in [[Tanzania]]
| taxon = Equus (Hippotigris)
| authority = C. H. Smith, 1841
| subdivision_ranks = Species
| subdivision =
†''[[Equus capensis|E. capensis]]''<br />
''[[Grévy's zebra|E. grevyi]]''<br />
†''[[Equus mauritanicus|E. mauritanicus]]''<br />
†''E. oldowayensis''<br />
''[[plains zebra|E. quagga]]''<br />
''[[mountain zebra|E. zebra]]''
| range_map = Zebra range.png
| range_map_caption = Modern range of the three living zebra species
}}

'''Zebras''' ({{IPAc-en|US|ˈ|z|iː|b|r|ə|z}}, {{IPAc-en|UK|ˈ|z|ɛ|b|r|ə|z|,_|ˈ|z|iː|-}})<ref>{{cite LPD|3}}</ref> (subgenus '''''Hippotigris''''') are African [[equines]] with distinctive black-and-white striped [[Animal coat|coat]]s. There are three [[Extant taxon|living]] species: [[Grévy's zebra]] (''Equus grevyi''), the [[plains zebra]] (''E. quagga''), and the [[mountain zebra]] (''E. zebra''). Zebras share the genus ''[[Equus (genus)|Equus]]'' with [[Wild horse|horses]] and [[Asinus|asses]], the three groups being the only living members of the family [[Equidae]]. Zebra stripes come in different patterns, unique to each individual. Several theories have been proposed for the function of these patterns, with most evidence supporting them as a deterrent for biting flies. Zebras inhabit [[East Africa|eastern]] and [[southern Africa]] and can be found in a variety of habitats such as [[savannahs]], [[grasslands]], woodlands, [[shrubland]]s, and mountainous areas.

Zebras are primarily [[grazing (behaviour)|grazers]] and can subsist on lower-quality vegetation. They are preyed on mainly by [[lion]]s, and typically flee when threatened but also bite and kick. Zebra species differ in [[Sociality|social behaviour]], with plains and mountain zebra living in stable [[Harem (zoology)|harems]] consisting of an adult male or [[stallion]], several adult females or [[mares]], and their young or [[foals]]; while Grévy's zebra live alone or in loosely associated herds. In harem-holding species, adult females mate only with their harem stallion, while male Grévy's zebras establish [[Territory (animal)|territories]] which attract females and the species is [[Polygynandry|polygynandrous]]. Zebras communicate with various vocalisations, body postures and facial expressions. [[Social grooming]] strengthens social bonds in plains and mountain zebras.

Zebras' dazzling stripes make them among the most recognisable mammals. They have been featured in art and stories in Africa and beyond. Historically, they have been highly sought by exotic animal collectors, but unlike [[horses]] and [[donkeys]], zebras have never been completely [[domesticated]]. The [[International Union for Conservation of Nature]] (IUCN) lists Grévy's zebra as [[Endangered species|endangered]], the mountain zebra as [[Vulnerable species|vulnerable]] and the plains zebra as [[Near-threatened species|near-threatened]]. The [[quagga]] (''E. quagga quagga''), a type of plains zebra, was driven to extinction in the 19th century. Nevertheless, zebras can be found in numerous protected areas.

==Etymology==
The English name "zebra" derives from [[Italian language|Italian]], [[Spanish language|Spanish]] or [[Portuguese language|Portuguese]].<ref name="etymology">{{cite web |title=Zebra |website=[[Online Etymology Dictionary]] |access-date=22 June 2020 |url=https://www.etymonline.com/word/zebra}}</ref><ref>{{cite web |title=Zebra |website=[[Lexico]] |access-date=25 June 2020 |url=https://www.lexico.com/en/definition/zebra|archive-url=https://web.archive.org/web/20200627194124/https://www.lexico.com/en/definition/zebra|archive-date=27 June 2020}}</ref> Its origins may lie in the [[Latin]] ''equiferus'', meaning "wild horse". ''Equiferus'' appears to have entered into Portuguese as ''ezebro'' or ''zebro'', which was originally used for a [[zebro|mysterious equine]] reported in the wilds of the [[Iberian Peninsula]] during the Middle Ages. In 1591, Italian explorer [[Filippo Pigafetta]] recorded "zebra" being used to refer to the African animals by Portuguese visitors to the continent.<ref name="zebro">{{Cite journal |doi=10.5252/az2015n1a2 |title=The Iberian Zebro: what kind of a beast was it? |journal=Anthropozoologica |volume=50 |year=2015 |last1=Nores |first1=Carlos |last2=Muñiz |first2=Arturo Morales |last3=Rodríguez |first3=Laura Llorente |last4=Bennett |first4=E. Andrew |last5=Geigl |first5=Eva-María|pages=21–32 |s2cid=55004515 |url=https://zenodo.org/record/4313384 |hdl=10651/34413 |hdl-access=free }}</ref> In ancient times, the zebra was called ''hippotigris'' ("horse tiger") by the [[Ancient Greece|Greeks]] and [[Ancient Rome|Romans]].<ref name="zebro" />{{sfn|Plumb|Shaw|2018|p=54}}

The word ''zebra'' was traditionally pronounced with a long initial vowel, but over the course of the 20th century the pronunciation with the short initial vowel became the norm in [[British English]].<ref>{{cite journal |last=Wells |first=John |title=Our Changing Pronunciation |journal=Transactions of the Yorkshire Dialect Society |year=1997 |volume=XIX|pages=42–48 |url=http://www.phon.ucl.ac.uk/home/wells/yorksdial-uni.htm |archive-url=https://web.archive.org/web/20141007035500/http://www.phon.ucl.ac.uk/home/wells/yorksdial-uni.htm |archive-date=2014-10-07 |url-status=live |access-date=2014-02-06}}</ref> The pronunciation with a long initial vowel remains standard in [[American English]].<ref>{{cite web |title=Zebra |url=https://dictionary.cambridge.org/pronunciation/english/zebra |website=[[Cambridge Advanced Learner's Dictionary|Cambridge Dictionary]] |access-date=26 May 2020}}</ref>

==Taxonomy==
{{Further|Evolution of the horse}}
Zebras are classified in the genus ''[[Equus (genus)|Equus]]'' (known as equines) along with [[wild horse|horses]] and [[Asinus|asses]]. These three groups are the only living members of the family [[Equidae]].<ref name="MacDonald" /> The [[plains zebra]] and [[mountain zebra]] were traditionally placed in the [[subgenus]] ''Hippotigris'' (C. H. Smith, 1841) in contrast to the [[Grévy's zebra]] which was considered the sole species of subgenus ''Dolichohippus'' (Heller, 1912).<ref name="Prothero 2003">{{Cite book |title=Horns, Tusks, and Flippers: The Evolution of Hoofed Mammals |last1=Prothero |first1=D. R. |last2=Schoch|first2= 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=216–218 |year=2003 |isbn=978-0-8018-7135-1}}</ref><ref>{{cite web|title=Hippotigris|website=ITIS|url=https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=926069#null|access-date=31 August 2020}}</ref><ref>{{cite web|title=Dolichohippus|website=ITIS|url=https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=926067#null|access-date=31 August 2020}}</ref> Groves and Bell (2004) placed all three species in the subgenus ''Hippotigris''.<ref name="GrovesBell2004">{{cite journal |author=Groves|first1= C. P. |last2=Bell|first2= 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> A 2013 [[phylogenetic]] study found that the plains zebra is more closely related to Grévy's zebras than mountain zebras.<ref name="Vilstrup">{{cite journal |author=Vilstrup|first1= Julia T. |last2=Seguin-Orlando|first2= A. |last3=Stiller|first3= M. |last4=Ginolhac|first4= A. |last5=Raghavan|first5= M. |last6=Nielsen|first6= S. C. A. |year=2013 |title=Mitochondrial phylogenomics of modern and ancient equids |journal=[[PLoS One]] |volume=8 |issue=2 |article-number=e55950 |doi=10.1371/journal.pone.0055950 |display-authors=etal |pmid=23437078 |pmc=3577844|bibcode= 2013PLoSO...855950V |doi-access= free }}</ref> The extinct [[quagga]] was originally classified as a distinct species.<ref>{{cite book |last1=Groves|first1= C. |last2=Grubb|first2= P. |year=2011 |title=Ungulate Taxonomy |publisher=Johns Hopkins University Press |page=16 |isbn=978-1-4214-0093-8}}</ref> Later genetic studies have placed it as the same species as the plains zebra, either a subspecies or just the southernmost population.<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><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> Molecular evidence supports zebras as a [[monophyletic]] [[Lineage (evolution)|lineage]].<ref name="Vilstrup" /><ref name="Forstén">{{cite journal |author=Forstén|first= Ann |year=1992 |title=Mitochondrial-DNA timetable and the evolution of ''Equus'': of molecular and paleontological evidence |journal=Annales Zoologici Fennici |volume=28 |pages=301–309 |url=http://www.sekj.org/PDF/anzf28/anz28-301-309.pdf}}</ref><ref>{{cite journal |last1=Ryder |first1=O. A. |last2=George |first2=M. |year=1986 |title=Mitochondrial DNA evolution in the genus ''Equus'' |journal=[[Molecular Biology and Evolution]] |volume=3 |issue=6 |pages=535–546 |doi=10.1093/oxfordjournals.molbev.a040414 |pmid=2832696 |url=http://mbe.library.arizona.edu/data/1986/0306/5geor.pdf |doi-access=free |access-date=13 November 2015 |archive-date=28 May 2016 |archive-url=https://web.archive.org/web/20160528170756/http://mbe.library.arizona.edu/data/1986/0306/5geor.pdf }}</ref>

''Equus'' originated in North America and direct [[paleogenomics|paleogenomic]] sequencing of a 700,000-year-old middle Pleistocene horse [[metapodial]] bone from Canada implies a date of 4.07 million years ago (mya) for the most [[recent common ancestor]] of the equines within a range of 4.0 to 4.5 mya.<ref name="ReferenceA">{{cite journal |last1=Orlando|first1= L.|last2 =Ginolhac|first2= A.|last3= Zhang|first3= G.|last4= Froese|first4= D.|last5= Albrechtsen|first5= A.|last6= Stiller|first6= M.|display-authors= etal |title=Recalibrating Equus evolution using the genome sequence of an early Middle Pleistocene horse |journal=[[Nature (journal)|Nature]] |volume=499 |issue=7456 |pages=74–78 |date=July 2013 |pmid=23803765 |doi=10.1038/nature12323|bibcode= 2013Natur.499...74O|s2cid= 4318227}}</ref> Horses [[Split (phylogenetics)|split]] from asses and zebras around this time and equines colonised Eurasia and Africa around 2.1–3.4 mya. Zebras and asses diverged from each other close to 2 mya. The mountain zebra diverged from the other species around 1.6 mya and the plains and Grévy's zebra split 1.4 mya.<ref name=Jónsson2014>{{cite journal|last1=Jónsson|first1=Hákon|last2=Schubert|first2=Mikkel|last3=Seguin-Orlando|first3=Andaine|last4=Orlando|first4=Ludovic|year=2014|title=Speciation with gene flow in equids despite extensive chromosomal plasticity|journal=Proceedings of the National Academy of Sciences|volume=111|issue=52|pages=18655–18660|doi=10.1073/pnas.1412627111 |pmid=25453089 |pmc=4284605 |bibcode=2014PNAS..11118655J |doi-access=free}}</ref>

A 2017 mitochondrial DNA study placed the Eurasian ''[[Equus ovodovi]]'' and the subgenus ''Sussemionus'' lineage as closer to zebras than to asses.<ref>{{Cite journal |last1=Druzhkova |first1=Anna S. |last2=Makunin |first2=Alexey I. |last3=Vorobieva |first3=Nadezhda V. |last4=Vasiliev |first4=Sergey K. |last5=Ovodov |first5=Nikolai D. |last6=Shunkov |first6=Mikhail V. |last7=Trifonov |first7=Vladimir A. |last8=Graphodatsky |first8=Alexander S. |date=January 2017 |title=Complete mitochondrial genome of an extinct ''Equus (Sussemionus) ovodovi'' specimen from Denisova cave (Altai, Russia) |journal=Mitochondrial DNA Part B |language=en |volume=2 |issue=1 |pages=79–81 |doi=10.1080/23802359.2017.1285209 |pmid=33473722 |pmc=7800821 |issn=2380-2359 |doi-access=free}}</ref> However, other studies disputed this placement, finding the ''Sussemionus'' lineage basal to the zebra+asses group, but suggested that the ''Sussemionus'' lineage may have received gene flow from zebras.<ref>{{Cite journal |last1=Cai |first1=Dawei |last2=Zhu |first2=Siqi |last3=Gong |first3=Mian |last4=Zhang |first4=Naifan |last5=Wen |first5=Jia |last6=Liang |first6=Qiyao |last7=Sun |first7=Weilu |last8=Shao |first8=Xinyue |last9=Guo |first9=Yaqi |last10=Cai |first10=Yudong |last11=Zheng |first11=Zhuqing |last12=Zhang |first12=Wei |last13=Hu |first13=Songmei |last14=Wang |first14=Xiaoyang |last15=Tian |first15=He |date=2022-05-11 |title=Radiocarbon and genomic evidence for the survival of Equus Sussemionus until the late Holocene |journal=eLife |language=en |volume=11 |article-number=e73346 |doi=10.7554/eLife.73346 |issn=2050-084X |pmc=9142152 |pmid=35543411 |doi-access=free }}</ref>

The cladogram of ''Equus'' below is based on Vilstrup and colleagues (2013) and Jónsson and colleagues (2014):<ref name="Vilstrup" /><ref name=Jónsson2014/>
{{clade| style=font-size:85%; line-height:90%;
|label1=''Equus''
|1={{clade
 |1={{clade
|label2=
  |1={{clade
 |label1='''Zebras'''
 |label2=[[Asinus|Wild asses]]
   |1={{clade
    |1=[[Mountain zebra]] (''E. zebra'') [[File:The book of the animal kingdom (Plate XVII) (white background).jpg|40 px]]
    |2={{clade
     |1=[[Plains zebra]] (''E. quagga'') [[File:NIE 1905 Horse - Burchell's zebra.jpg|40 px]]
     |2=[[Grévy's zebra]] (''E. grevyi'') [[File:Equus grevyi (white background).png|40 px]]}} }}
   |2={{clade
    |1={{clade
     |1=[[Kiang]] (''E. kiang'')  [[File:Equus hemionus - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|40 px]]
     |2=[[Onager]] (''E. hemionus'') [[File:Hémippe (white background).jpg|40 px]]}}
    |2=[[African wild ass]] (''E. africanus'') [[File:Âne d'Ethiopie (white background).jpg|40 px]]}} }} }}
  |label2=[[Wild horse|Horses]]
 |2={{clade
    |1=[[Horse]] (''E. ferus caballus'') [[File:NIEdot332 white background 2.jpg|40 px]]
    |2=[[Przewalski's horse]] (''E. ferus przewalski'') [[File:The Soviet Union 1959 CPA 2325 stamp (Przewalski's Horse) white background.jpg|40 px]]}} }} }}

===Extant species===
{| class="wikitable"
|+ style="text-align: centre;" |
! Name
! Dimensions
! Description
! Distribution
! Subspecies
! Chromosomes
! Image
|-
| [[Grévy's zebra]] (''Equus grevyi'')
|'''Body length''': {{cvt|250|–|300|cm}}<br />'''Tail length''': {{cvt|38|–|75|cm}}<br />'''Shoulder height''': {{cvt|125|–|160|cm}}<br />'''Weight''': {{cvt|352|–|450|kg}}.{{sfn|Caro|2016|p=9}}
|Thin, elongated skull, robust neck and conical ears; narrow striping pattern with [[Concentric objects|concentric]] rump stripes, white belly and tail base and white line around the ashy muzzle.<ref name="MacDonald">{{Cite book |last=Rubenstein|first= D. I. |contribution=Horse, Zebras and Asses |year=2001 |title=The Encyclopedia of Mammals |edition=2nd |editor=MacDonald|editor-first= D. W. |publisher=[[Oxford University Press]] |pages=468–473 |isbn=978-0-7607-1969-5}}</ref><ref name="Churcher 1993" />{{sfn|Caro|2016|p=15}}
| [[East Africa|Eastern Africa]] including [[Horn of Africa|the Horn]];<ref name="Churcher 1993">{{cite journal |author=Churcher|first= C. S. |year=1993 |title=Equus grevyi |url=http://www.science.smith.edu/departments/Biology/VHAYSSEN/msi/pdf/i0076-3519-453-01-0001.pdf |journal=[[Mammalian Species]] |issue=453 |pages=1–9 |doi=10.2307/3504222 |jstor=3504222}}</ref> arid and semiarid [[grassland]]s and [[shrubland]]s.{{sfn|Caro|2016|p=14}}
|[[Monotypic taxon|Monotypic]]<ref name="Churcher 1993"/>
|46{{sfn|Caro|2016|p=14}}
|[[File:Grevy's Zebra Stallion.jpg|120px]]
|-
|[[Plains zebra]] (''Equus quagga'')
|'''Body length''': {{cvt|217|–|246|cm}}<br />'''Tail length''': {{cvt|47|–|56|cm|in}}<br />'''Shoulder height''': {{cvt|110|–|145|cm}}<br />'''Weight''': {{cvt|175|–|385|kg}}.{{sfn|Caro|2016|p=9}}
|Thick bodied with relatively short legs and an obtusely shaped skull profile with a protruding forehead and a more recessed nose area;<ref name="MacDonald" /><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> broad stripes, horizontal on the rump, with northern populations having more extensive striping while populations further south have whiter legs and bellies and more brown "shadow" stripes while the snout is black.<ref name="MacDonald" /><ref name="Estes 1991" />{{sfn|Caro|2016|pp=12–13}}<ref name="Skinner"/>
|Eastern and [[southern Africa]]; [[savannah]]s, grasslands and open woodlands.{{sfn|Caro|2016|p=11}}
|6<ref name="GrovesBell2004"/> or monotypic<ref name="PedersenAlbrechtsen2018"/>
|44{{sfn|Caro|2016|p=13}}
|[[File:Equus quagga burchellii - Etosha, 2014.jpg|120px]]
|-
|[[Mountain zebra]] (''Equus zebra'')
|'''Body length''': {{cvt|210|–|260|cm}}<br />'''Tail length''': {{cvt|40|–|55|cm}}<br />'''Shoulder height''': {{cvt|116|–|146|cm}}<br />'''Weight''': {{cvt|204|–|430|kg}}.{{sfn|Caro|2016|p=9}}
|[[Eye sockets]] more circular and positioned farther back, a squarer [[Nuchal lines|nuchal crest]], [[dewlap]] present under neck and compact hooves; stripes intermediate in width between the other species, with gridiron and horizontal stripes on the rump, while the belly is white and the black muzzle is lined with [[Chestnut (color)|chestnut]] or orange.<ref name="Penzhorn 1988">{{cite journal |last1=Penzhorn |first1=B. L. |year=1988 |title=Equus zebra |journal=Mammalian Species |issue=314 |pages=1–7 |doi=10.2307/3504156|jstor=3504156 |s2cid=253987177 }}</ref><ref name="MacDonald" /><ref name="handbook" />{{sfn|Caro|2016|p=14}}
|Southwestern Africa; mountains, rocky uplands and [[Karoo]] shrubland.{{sfn|Caro|2016|p=11}}<ref name="Penzhorn 1988" /><ref name="Skinner"/>
|2<ref name="Penzhorn 1988" />
|32{{sfn|Caro|2016|p=14}}
|[[File:Equus zebra hartmannae - Etosha 2015.jpg|120px]]
|}

===Fossil record===
[[File:Equus mauritanicus.JPG|thumb|right|alt= A fossil skull of ''Equus mauritanicu'' | Fossil skull of ''Equus mauritanicus'']]
In addition to the three living species, some fossil zebras and relatives have also been identified. ''E. oldowayensis'' is identified from remains in Olduvai Gorge dating to 1.8 mya.<ref name="Churcher2006" /> Fossil skulls of ''E. mauritanicus'' from Algeria which date to around 1 mya appears to show affinities with the plains zebra.<ref name="Azzaroli">{{Cite journal |last1=Azzaroli |first1=A. |last2=Stanyon |first2=R. |doi=10.1007/BF03001000 |title=Specific identity and taxonomic position of the extinct Quagga |journal=Rendiconti Lincei |volume=2 |issue=4 |page=425 |year=1991 |s2cid=87344101 }}</ref><ref>{{cite journal|author=Eisenmann|first= V.|year=2008|title=Pliocene and Pleistocene equids: palaeontology versus molecular biology|journal= Courier Forschungsinstitut Senckenberg|volume=256|pages=71–89|url=https://www.researchgate.net/publication/281338907}}</ref> ''[[Equus capensis|E. capensis]]'', known as the Cape zebra, appeared around 2 mya and lived throughout southern and eastern Africa.<ref name="Badenhorst2019">{{cite journal |author=Badenhorst|first1= S. |last2=Steininger|first2= C. M. |year=2019 |title=The Equidae from Cooper's D, an early Pleistocene fossil locality in Gauteng, South Africa |journal=[[PeerJ]] |volume=7 |article-number= e6909 |doi=10.7717/peerj.6909 |pmid= 31143541 |pmc= 6525595 |doi-access=free}}</ref><ref name="Churcher2006">{{cite journal |author=Churcher|first= C. S. |year=2006 |title=Distribution and history of the Cape zebra (''Equus capensis'') in the Quarternary of Africa |journal=[[Transactions of the Royal Society of South Africa]] |volume=61 |issue=2 |pages=89–95 |doi=10.1080/00359190609519957|bibcode= 2006TRSSA..61...89C |s2cid= 84203907 }}</ref>

===Hybridisation===
{{Main|Zebroid}}
Fertile hybrids have been reported in the wild between plains and Grévy's zebra.<ref>{{cite journal |first1=J. E.|last1= Cordingley |first2=S. R.|last2= Sundaresan |first3=I. R.|last3= Fischhoff |first4=B.|last4= Shapiro |first5=J.|last5= Ruskey |first6=D. I.|last6= 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> Hybridisation has also been recorded between the plains and mountain zebra, though it is possible that these are infertile due to the difference in chromosome numbers between the two species.<ref>{{cite book |author=Giel|first1= E.-M. |last2=Bar-David|first2= S. |last3=Beja-Pereira|first3= A. |last4=Cothern|first4= E. G. |last5=Giulotto|first5= E. |last6=Hrabar|first6= H. |last7=Oyunsuren|first7= T. |last8=Pruvost|first8= M. |year=2016 |contribution=Genetics and Paleogenetics of Equids |title=Wild Equids: Ecology, Management, and Conservation |editor=Ransom|editor-first= J. I. |editor2=Kaczensky|editor-first2=P. |publisher=Johns Hopkins University Press |page=99 |isbn=978-1-4214-1909-1}}</ref> Captive zebras have been bred with horses and [[donkeys]]; these are known as [[zebroids]]. A zorse is a cross between a zebra and a horse; a zonkey, between a zebra and a donkey; and a zoni, between a zebra and a [[pony]]. Zebroids are often born sterile with [[dwarfism]].<ref>{{cite web |author=Bittel|first= Jason |date=19 June 2015 |title=Hold Your Zorses: The sad truth about animal hybrids |publisher=[[Slate.com]] |access-date=16 May 2020 |url=https://slate.com/technology/2015/06/zonkeys-ligers-the-sad-truth-about-animal-hybrids.html}}</ref>

<gallery mode="packed" heights="150px">
Grévy's × Plains Zebra, ol pejeta imported from iNaturalist photo 401473965 (cropped).jpg|Grévy's × plains zebra hybrid, alongside plains zebras
Donkey × Plains Zebra, Cedarberg Guest Farm, Cedarville, South Africa imported from iNaturalist photo 11786274 (cropped).jpg|Donkey × plains zebra hybrid, in South Africa
</gallery>

==Characteristics==
{{Further|Equine anatomy}}
[[File:Equus grevyi 01.JPG|thumb|right|alt=Mounted skeleton of a Grévy's zebra Cranium, complete skeleton, left forefoot frontal, left forefoot side |Skeleton of a Grévy's zebra at the [[State Museum of Natural History Karlsruhe]]]]
As with all wild equines, zebras have barrel-chested bodies with [[Tail (horse)|tufted tails]], elongated faces and long necks with long, erect [[Mane (horse)|manes]]. Their thin legs are each supported by a spade-shaped toe covered in a hard [[horse hoof|hoof]]. Their [[horse teeth|dentition]] is adapted for [[grazing (behaviour)|grazing]]; they have large incisors that clip grass blades and rough molars and premolars well suited for grinding. Males have spade-shaped canines, which can be used as weapons in fighting. The eyes of zebras are at the sides and far up the head, which allows them to look over the tall grass while feeding. Their moderately long, erect ears are movable and can locate the source of a sound.<ref name="MacDonald" /><ref name="Estes 1991">{{Cite book |title=The Behavior Guide to African Mammals |author=Estes|first= R. |publisher=[[University of California Press]] |pages=235–248 |year=1991 |isbn=978-0-520-08085-0 |url=https://books.google.com/books?id=g977LsZHpcsC}}</ref><ref name="handbook">{{Cite book |last=Rubenstein|first= D. I. |contribution=Family Equidae: Horses and relatives |year=2011 |title=Handbook of the Mammals of the World|volume= 2: Hoofed Mammals |edition=1st |editor=Wilson|editor-first= D. E.|editor2= Mittermeier|editor-first2= R. A. |editor3= Llobet|editor-first3= T. |publisher=[[Lynx Edicions]] |pages=106–111 |isbn=978-84-96553-77-4}}</ref>

Unlike horses, zebras and asses have [[Chestnut (horse anatomy)|chestnut callosities]] present only on their front legs. In contrast to other living equines, zebras have longer front legs than back legs.<ref name="handbook" /> Diagnostic traits of the zebra skull include: its relatively small size with a straight dorsal outline, protruding eye sockets, narrower rostrum, less conspicuous [[postorbital bar]], separation of the [[Glossary of mammalian dental topography|metaconid]] and [[Glossary of mammalian dental topography|metastylid]] of the tooth by a V-shaped canal and rounded [[Tooth enamel|enamel wall]].<ref>{{cite journal |author=Badam|first1=  G. L. |last2=Tewari|first2=  B. S. |year=1974 |title=On the zebrine affinities of the Pleistocene horse ''Equus sivalensis'', ''falconer'' and ''cautley'' |journal=Bulletin of the Deccan College Post-Graduate and Research Institute |volume=34 |issue=1/4 |pages=7–11 |jstor=42931011}}</ref>

===Stripes===
{{Redirect|Zebra stripes||Zebra stripes (disambiguation)}}
[[File:Zebra species (ENG).png|thumb|left|alt=An illustration showing the three living zebra species |Comparative illustration of living zebra species]]
Zebras are easily recognised by their bold black-and-white striping patterns. The [[Animal coat|coat]] appears to be white with black stripes, as indicated by the belly and legs when unstriped, but the skin is black.<ref name="Bard1977" /><ref>{{cite web |author=Langley|first=  Liz |date=4 March 2017 |title=Do Zebras Have Stripes On Their Skin? |work=National Geographic |access-date=2 June 2020 |url=https://www.nationalgeographic.com/news/2017/03/animals-skin-colors-zebras-big-cats/|archive-url=https://web.archive.org/web/20200801013454/https://www.nationalgeographic.com/news/2017/03/animals-skin-colors-zebras-big-cats/|archive-date=1 August 2020}}</ref>{{sfn|Caro|2016|pp=14–15}} Young or [[foal]]s are born with brown and white coats, and the brown darkens with age.<ref name="Grub 1981" /><ref name="Churcher 1993" /> A [[Primitive_markings#Dorsal_stripe|dorsal stripe]] acts as the backbone for vertical stripes along the sides, from the head to the rump. On the snout they curve toward the nostrils, while the stripes above the front legs split into two branches. On the rump, they develop into species-specific patterns. The stripes on the legs, ears and tail are separate and horizontal.<ref name="Bard1977" />

Striping patterns are unique to an individual and heritable.{{sfn|Caro|2016|pp=7, 19}} During [[embryonic development]], the stripes appear at eight months, but the patterns may be determined at three to five weeks. For each species there is a point in embryonic development where the stripes are perpendicular to the dorsal line and spaced {{convert|0.4|mm|in|abbr=on}} apart. However, this happens at three weeks of development for the plains zebra, four weeks for the mountain zebra, and five for Grévy's zebra. The difference in timing is thought to be responsible for the differences in the striping patterns of the different species.<ref name="Bard1977">{{cite journal |author=Bard|first=  J. |year=1977 |title=A unity underlying the different zebra patterns |journal=[[Journal of Zoology]] |volume=183 |issue=4 |pages=527–539 |doi=10.1111/j.1469-7998.1977.tb04204.x}}</ref>

Various abnormalities of the patterns have been documented in plains zebras. In "[[melanistic]]" zebras, dark stripes are highly concentrated on the torso but the legs are whiter. "Spotted" individuals have broken up black stripes around the dorsal area.<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.{{sfn|Caro|2016|p=20}} 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/ |archive-date=29 March 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>

====Function====
The function of stripes in zebras has been discussed among biologists since at least the 19th century.{{sfn|Caro|2016|p=1}} Popular hypotheses include the following:
* The '''[[crypsis]] hypothesis''' suggests that the stripes allow the animal to blend in with its environment or [[disruptive coloration|break up its outline]]. This was the earliest hypothesis and proponents argued that the stripes were particularly suited for camouflage in tall grassland and woodland habitat. [[Alfred Wallace]] also wrote in 1896 that stripes make zebras less noticeable at night. Biologist [[Tim Caro]] notes that zebras graze in open habitat and do not behave cryptically, being noisy, fast, and social and do not freeze when a predator is near. In addition, the camouflaging stripes of woodland living ungulates like [[Bongo (antelope)|bongos]] and [[Cape bushbuck|bushbucks]] are much less vivid with less contrast with the background colour.{{sfn|Caro|2016|pp=2–3, 23, 48, 50}} A 1987 [[Fourier analysis]] study concluded that the [[spatial frequencies]] of zebra stripes do not line up with their environment,<ref>{{cite journal |last1=Godfrey|first1=  D. |last2=Lythgoe|first2=  J. N. |last3=Rumball|first3=  D. A. |year=1987 |title=Zebra stripes and tiger stripes: the spatial frequency distribution of the pattern compared to that of the background is significant in display and crypsis |journal=[[Biological Journal of the Linnean Society]] |volume=32 |issue=4 |pages=427–433 |doi=10.1111/j.1095-8312.1987.tb00442.x}}</ref> while a 2014 study of wild equine species and subspecies could not find any correlations between striping patterns and woodland habitats.<ref name="Caro" /> Melin and colleagues (2016) found that [[lion]]s and hyenas do not appear to perceive the stripes when they are a certain distance away at daytime or nighttime, thus making the stripes useless in blending in except when the predators are close enough by which they could smell or hear their target. They also found that the stripes do not make the zebra less noticeable than solidly coloured herbivores on the open plains. They suggested that stripes may give zebras an advantage in woodlands, as the dark stripes could line up with the outlines of tree branches and other vegetation.<ref>{{cite journal|last1=Melin|first1=A. D.|last2=Kline|first2=D. W.|last3=Hiramatsu|first3=C|last4=Caro|first4=T|year=2016|title=Zebra stripes through the eyes of their predators, zebras, and humans|journal=PLOS ONE|volume=11|issue=1|article-number=e0145679|doi=10.1371/journal.pone.0145679|doi-access=free|pmid=26799935 |pmc=4723339|bibcode=2016PLoSO..1145679M }}</ref>
[[File:Mountain zebra stripes.jpg|thumb|right|alt=Closeup shot of mountain zebra stripes |Closeup of mountain zebra stripes]]
* The '''confusion hypothesis''' states that the stripes confuse predators, be it by: making it harder to distinguish individuals in a group as well as determining the number of zebras in a group; making it difficult to determine an individual's outline when the group runs away; reducing a predator's ability to keep track of a target during a chase; [[dazzle camouflage|dazzling]] an assailant so they have difficulty making contact; or making it difficult for a predator to deduce the zebra's size, speed and direction via [[motion dazzle]]. This theory has been proposed by several biologists since at least the 1970s.{{sfn|Caro|2016|pp=72–81, 86}} A 2014 computer study of zebra stripes found that they may create a [[wagon-wheel effect]] and/or [[barber pole illusion]] when in motion. The researchers concluded that this could be used against mammalian predators or biting flies.<ref name="How2014">{{cite journal |author=How|first1=  M. J. |last2=Zanker|first2=  J. M. |year=2014 |title=Motion camouflage induced by zebra stripes |journal=Zoology |volume=117 |issue=3 |pages=163–170 |doi=10.1016/j.zool.2013.10.004|pmid= 24368147 |bibcode= 2014Zool..117..163H }}</ref> The use of the stripes for confusing mammalian predators has been questioned. Caro suggests that the stripes of zebras could make groups seem smaller, and thus more likely to be attacked. Zebras also tend to scatter when fleeing from attackers and thus the stripes could not break up an individual's outline. Lions, in particular, appear to have no difficulty targeting and catching zebras when they get close and take them by ambush.{{sfn|Caro|2016|pp=80, 92}} In addition, no correlations have been found between the number of stripes and populations of mammal predators.<ref name="Caro" /> Hughes and colleagues (2021) disputed the idea of motion dazzle and concluded that moving objects that are solidly grey or have less contrasted patterns are actually more likely to escape being caught.<ref>{{cite journal|last1=Hughes|first1=A. E.|last2=Griffiths|first2=D|last3=Troscianko|first3=J|last4=Kelley|first4=L. A.|year=2021|title=The evolution of patterning during movement in a large-scale citizen science game|journal=Proceedings of the Royal Society B: Biological Sciences|volume=288|issue=1942 |article-number=20202823|doi=10.1098/rspb.2020.2823|pmid=33434457 |pmc=7892415 }}</ref> 
* The '''[[Aposematism|aposematic]] hypothesis''' suggests that the stripes serve as warning colouration. This hypothesis was first suggested by Wallace in 1867 and discussed in more detail by [[Edward Bagnall Poulton]] in 1890. As with known aposematic mammals, zebras are recognizable up close, live in more open environments, have a high risk of predation and do not hide or act inconspicuous. However, Caro notes that stripes do not work on lions because they frequently prey on zebras, though they may work on smaller predators, and zebras are not slow-moving enough to need to ward off threats. In addition, zebras do not possess adequate defenses to back up the warning pattern.{{sfn|Caro|2016|pp=55, 57–58, 68}}
* The '''social function hypothesis''' states that stripes serve a role in intraspecific or individual recognition, social bonding, mutual grooming or a signal of [[Fitness (biology)|fitness]]. [[Charles Darwin]] wrote in 1871{{efn|The cited source cites the 1896 edition of Darwin's 1871 book ''[[The Descent of Man, and Selection in Relation to Sex]]''.}} that "a female zebra would not admit the addresses of a male ass until he was painted so as to resemble a zebra" while Wallace stated in 1871 that: "The stripes therefore may be of use by enabling stragglers to distinguish their fellows at a distance". Regarding species and individual identification, Caro notes that zebra species have limited range overlap with each other and horses can recognise each other using visual communication.{{sfn|Caro|2016|pp=6–7, 139–148, 150}} In addition, no correlation has been found between striping and social behaviour or group numbers among equines,<ref name="Caro" /> and no link has been found between fitness and striping.{{sfn|Caro|2016|p=150}}
* The '''[[thermoregulation|thermoregulatory]] hypothesis''' suggests that stripes help to control a zebra's body temperature. In 1971, biologist H. A. Baldwin noted that heat would be absorbed by the black stripes and reflected by the white ones and in 1990, zoologist [[Desmond Morris]] suggested that the stripes create cooling [[convection current]]s.{{sfn|Caro|2016|p=7}} A 2019 study supported this, finding that where the faster air currents of the warmer black stripes meet those of the white, [[Vortex|air swirl]]s form. The researchers also concluded that during the hottest times of the day, zebras [[Hackles|erect]] their black hair to release heat from the skin and flatten it again when it gets cooler.<ref>{{cite journal |last1=Cobb|first1=  A. |last2=Cobb|first2=  S. |year=2019 |title=Do zebra stripes influence thermoregulation? |journal=[[Journal of Natural History]] |volume=53 |issue=13–14 |pages=863–879 |doi=10.1080/00222933.2019.1607600|bibcode= 2019JNatH..53..863C |s2cid= 196657566 }}</ref> Larison and colleagues (2015) determined that environmental temperature is a strong predictor for zebra striping patterns.<ref>{{cite journal |last1=Larison |first1=Brenda |last2=Harrigan |first2=Ryan J. |last3=Thomassen |first3=Henri A. |last4=Rubenstein |first4=Daniel I. |last5=Chan-Golston |first5=Alec M. |last6=Li |first6=Elizabeth |last7=Smith |first7=Thomas B. |year=2015 |title=How the zebra got its stripes: a problem with too many solutions |url=http://rsos.royalsocietypublishing.org/content/2/1/140452tempa |journal=[[Royal Society Open Science]] |volume=2 |issue=1 |article-number=140452 |doi=10.1098/rsos.140452 |pmid=26064590 |pmc=4448797|bibcode=2015RSOS....240452L }}</ref> Others have found no evidence that zebras have lower body temperatures than other ungulates whose habitat they share, or that striping correlates with temperature.{{sfn|Caro|2016|pp=158–161}}<ref name="Caro" /> A 2018 experimental study which dressed water-filled metal barrels in horse, zebra and cattle hides concluded that the zebra stripes had no effect on thermoregulation.<ref>{{cite journal|last1=Horváth |first1=Gábor |last2=Pereszlényi |first2=Ádám |last3=Száz |first3= Dénes|last4=Barta |first4= András |last5= Jánosi |first5= Imre M. |last6=Gerics |first6= Balázs |last7=Åkesson |first7=Susanne|year=2018|title=Experimental evidence that stripes do not cool zebras|journal=Scientific Reports|volume=8|issue=1|page=9351|doi=10.1038/s41598-018-27637-1|pmid=29921931 |pmc=6008466 |bibcode=2018NatSR...8.9351H |doi-access=free }}</ref>
[[File:Journal.pone.0210831.g001.png|thumb|right|alt=Comparison of horse fly flight patterns on horses and zebras |Comparison of flight patterns and contact/landings of horse flies around domestic horses (a-c) and plains zebras (d-f).<ref name="Caro2" />]]
* The '''fly protection hypothesis''' holds that the stripes deter [[blood-sucking]] flies. [[Horse fly|Horse flies]], in particular, spread diseases that are lethal to equines such as [[African horse sickness]], [[equine influenza]], [[equine infectious anemia]] and [[trypanosomiasis]]. In addition, zebra hair is about as long as the mouthparts of these flies.<ref name="Caro" /> This hypothesis is the most strongly supported by the evidence.<ref name="Caro2" /><ref name=Tombak2022/> It was found that flies preferred landing on solidly coloured surfaces over those with black-and-white striped patterns in 1930 by biologist R. Harris,{{sfn|Caro|2016|p=5}} and this was proposed to have been a function of zebra stripes in a 1981 study.<ref>{{cite journal|last1=Waage|first1=J. K.|year=1981|title=How the zebra got its stripes - biting flies as selective agents in the evolution of zebra coloration|journal=Journal of the Entomological Society of Southern Africa|volume=44|issue=2|pages=351–358|hdl=10520/AJA00128789_3800}}</ref> A 2014 study found a correlation between striping and overlap with horse and [[tsetse fly]] populations and activity.<ref name="Caro">{{cite journal |last1=Caro|first1=  T. |last2=Izzo|first2=  A. |last3=Reiner|first3=  R. C. |last4=Walker|first4=  H. |last5=Stankowich|first5=  T. |year=2014 |title=The function of zebra stripes |journal=[[Nature Communications]] |volume=5 |article-number=3535 |bibcode= 2014NatCo...5.3535C|doi=10.1038/ncomms4535 |pmid=24691390 |doi-access=free}}</ref> Other studies have found that zebras are rarely targeted by these insect species.{{sfn|Caro|2016|pp=196–197}} Caro and colleagues (2019) studied captive zebras and horses and observed that neither could deter flies from a distance, but zebra stripes kept flies from landing, both on zebras and horses dressed in zebra [[Animal print|print]] coats.<ref name="Caro2">{{cite journal |author=Caro|first1= T. |last2=Argueta|first2= Y. |last3=Briolat|first3= E. S. |last4=Bruggink|first4= J. |last5=Kasprowsky|first5= M. |last6=Lake|first6= J. |last7=Mitchell|first7= M. |last8=Richardson|first8= S. |last9=How|first9= M. |year=2019 |title=Benefits of zebra stripes: behaviour of tabanid flies around zebras and horses |journal=PLOS ONE |volume=14 |issue=2 |article-number=e0210831 |doi=10.1371/journal.pone.0210831 |pmid= 30785882 |pmc= 6382098 |bibcode= 2019PLoSO..1410831C |doi-access=free}}</ref> There does not appear to be any difference in the effectiveness of repelling flies between the different zebra species; thus the difference in striping patterns may have evolved for other reasons.<ref name=Tombak2022>{{cite journal|last1=Tombak|first1=K. J.|last2=Gersick|first2=A. S.|last3=Reisinger|first3=L. V.|last4=Larison|first4=B|last5=Rubenstein|first5=D. I.|year=2022|title=Zebras of all stripes repel biting flies at close range|journal=Scientific Reports |volume=22|issue=18617|page=18617 |doi=10.1038/s41598-022-22333-7|pmid=36329147 |pmc=9633588 |bibcode=2022NatSR..1218617T }}</ref> White or light stripes painted on dark bodies have also been found to reduce fly irritations in both cattle and humans.<ref>{{cite journal |last1=Kojima|first1= T. |last2=Oishi|first2= K. |last3=Matsubara|first3= Y. |last4=Uchiyama|first4= Y. |last5=Fukushima|first5= Y. |year=2020 |title=Cows painted with zebra-like striping can avoid biting fly attack |journal=PLOS ONE |volume=15 |issue=3 |article-number=e0231183 |doi=10.1371/journal.pone.0231183 |pmid= 32214400 |pmc= 7098620 |doi-access=free}}</ref><ref>{{cite journal |last1=Horváth|first1= G. |last2=Pereszlényi|first2= Á. |last3=Åkesson|first3= S. |last4=Kriska|first4= G. |year=2019 |title=Striped bodypainting protects against horseflies |journal=Royal Society Open Science |volume=6 |issue=1 |article-number= 181325 |doi=10.1098/rsos.181325 |pmid= 30800379 |pmc= 6366178 |bibcode= 2019RSOS....681325H |doi-access=free}}</ref> How the stripes repel flies is less clear.<ref name=Tombak2022/> A 2012 study concluded that they disrupt the [[Polarization (physics)|polarised]] light patterns these insects use to locate water and habitat,<ref>{{cite journal |last1=Egri |first1=Ádám |last2=Blahó |first2=Miklós |last3=Kriska |first3=György |last4=Farkas |first4=Róbert |last5=Gyurkovszky |first5=Mónika |last6=Åkesson |first6=Susanne |last7=Horváth |first7=Gábor |year=2012 |title=Polarotactic tabanids find striped patterns with brightness and/or polarization modulation least attractive: an advantage of zebra stripes |journal=[[Journal of Experimental Biology]] |volume=215 |issue=5 |pages=736–745 |doi=10.1242/jeb.065540 |pmid=22323196 |doi-access=free|bibcode=2012JExpB.215..736E }}</ref> though subsequent studies have refuted this.<ref name=Caro2023/><ref name=Takács2022/><ref>{{cite journal|last1=Britten|first1=K. H.|last2=Thatcher|first2=T. D.|last3=Caro|first3=T|year=2016|title=Zebras and biting flies: quantitative analysis of reflected light from zebra coats in Their natural habitat|journal=PLOS ONE|volume=11|issue=5|article-number=e0154504|doi=10.1371/journal.pone.0154504|pmid=27223616 |pmc=4880349 |bibcode=2016PLoSO..1154504B |doi-access=free }}</ref> Stripes do not appear to work like a barber pole against flies since [[Check (pattern)|checkered pattern]]s also repel them.<ref name=Caro2023/><ref>{{cite journal|last1=How|first1=M. J.|last2=Gonzales|first2=D.|last3=Irwin|first3=A.|last4=Caro|first4=T.|year=2020|title=Zebra stripes, tabanid biting flies and the aperture effect|journal=Proceedings of the Royal Society B: Biological Sciences|volume=287|issue=1933|article-number=20201521 |doi=10.1098/rspb.2020.1521|pmid=32811316|pmc=7482270}}</ref> There is also little evidence that zebra stripes confuse the insects via visual distortion or [[aliasing]].<ref name=Caro2023/> Takács and colleagues (2022) suggest that, when the animal is in sunlight, temperature gradients between the warmer dark stripes and cooler white stripes prevent horseflies from detecting the warm blood vessels underneath.<ref name=Takács2022>{{cite journal|last1=Takács|first1=P|last2=Száz|first2=D|last3=Vincze|first3=M|last4=Slíz-Balogh|first4=J|last5=Horváth|first5=G|year=2022|title=Sunlit zebra stripes may confuse the thermal perception of blood vessels causing the visual unattractiveness of zebras to horseflies|journal=Scientific Reports|volume=12|issue=10871|page=10871|doi=10.1038/s41598-022-14619-7|pmid=35927437|pmc=9352684|bibcode=2022NatSR..1210871T}}</ref> Caro and colleagues (2023) conclude that the high colour contrast and relative thinness of the patterns make it difficult for the insects to find a place to land.<ref name=Caro2023>{{cite journal|last1=Caro|first1=T|last2=Fogg|first2=E|last3=Stephens-Collins|first3=T|last4=Santon|first4=M|last5=How|first5=M. J.|year=2023|title=Why don't horseflies land on zebras?|journal=Journal of Experimental Biology|volume=226|issue=4|page=jeb244778|doi=10.1242/jeb.244778|pmid=36700395|pmc=10088525|bibcode=2023JExpB.226B4778C|s2cid=256273744}}</ref>

==Behaviour and ecology==
[[File:Dust bathing mountain zebra, Namibia.jpg|thumb|right|alt=Mountain zebra dust bathing |Mountain zebra dustbathing in [[Namibia]]]]
Zebras may travel or [[animal migration|migrate]] to wetter areas during the dry season.<ref name="Grub 1981" /><ref name="Estes 1991" /> Plains zebras have been recorded travelling {{cvt|500|km|mi}} between Namibia and Botswana, the longest land migration of mammals in Africa.<ref>{{cite journal |last1=Naidoo|first1= R. |last2=Chase|first2= M. J. |last3=Beytall|first3= P. |last4=Du Preez|first4= P. |year=2016 |title=A newly discovered wildlife migration in Namibia and Botswana is the longest in Africa |journal=Oryx |volume=50 |issue=1 |pages=138–146 |doi=10.1017/S0030605314000222|doi-access=free }}</ref> When migrating, they appear to rely on some memory of the locations where foraging conditions were best and may predict conditions months after their arrival.<ref>{{cite journal |author=Bracis|first1= C. |last2=Mueller|first2= 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 |article-number=20170449 |doi=10.1098/rspb.2017.0449 |pmid=28539516 |pmc=5454266}}</ref> Plains zebras are more water-dependent and live in [[Mesic habitat|moister]] environments than other species. They usually can be found {{cvt|10|–|12|km}} from a water source.<ref name="Grub 1981" /><ref name="Estes 1991" /><ref name="Skinner" /> Grévy's zebras can survive almost a week without water but will drink it every day when given the chance, and their bodies maintain water better than cattle.<ref>{{cite journal |author=Youth|first= H. |title=Thin stripes on a thin line |url=http://nationalzoo.si.edu/Publications/ZooGoer/2004/6/grevys.cfm |journal=Zoogoer |volume=33 |date=November–December 2004 |archive-date=26 October 2005  |archive-url=https://web.archive.org/web/20051026202556/http://nationalzoo.si.edu/Publications/ZooGoer/2004/6/grevys.cfm}}</ref><ref name="Churcher 1993" /> Mountain zebras can be found at elevations of up to {{cvt|2000|m}}.<ref>{{cite book |author=Woodward|first= Susan L. |year=2008 |title=Grassland Biomes |publisher=[[Greenwood Press]] |page=49 |isbn=978-0-313-33999-8}}</ref> Zebras sleep up to seven hours a day, standing up during the day and lying down during the night. They regularly use various objects as rubbing posts and will [[dust bathing|roll on the ground]].<ref name="Estes 1991" />
[[File:Cebras de Burchell (Equus quagga burchellii), vista aérea del delta del Okavango, Botsuana, 2018-08-01, DD 30.jpg|thumb|left|alt=Plains zebras drinking at a river |Plains zebras at [[Okavango Delta]], [[Botswana]]]]
A zebra's diet is mostly [[grass]]es and [[sedge]]s, but they will opportunistically consume [[bark (botany)|bark]], leaves, buds, fruits, and roots. Compared to [[Ruminantia|ruminants]], zebras have a simpler and less efficient digestive system. Nevertheless, they can subsist on lower-quality vegetation. Zebras may spend 60–80% of their time feeding, depending on the availability of vegetation.<ref name="MacDonald" /><ref name="Estes 1991" /> The plains zebra is a pioneer grazer, mowing down the upper, less nutritious grass canopy and preparing the way for more specialised grazers like [[wildebeest]], which depend on shorter and more nutritious grasses below.<ref>{{cite book |last1=Pastor|first1= J.|last2= Cohen|first2= U.|last3= Hobbs|first3= T. |year=2006 |contribution=The roles of large herbivores in ecosystem nutrient cycles |editor=Danell|editor-first= K. |title=Large Herbivore Ecology, Ecosystem Dynamics and Conservation |url=https://archive.org/details/largeherbivoreec00dane_862 |url-access=limited |publisher=[[Cambridge University Press]] |page=[https://archive.org/details/largeherbivoreec00dane_862/page/n312 295] |isbn=978-0-521-53687-5}}</ref>

Zebras are preyed on mainly by lions. [[Leopards]], [[cheetahs]], [[spotted hyenas]], [[brown hyena]]s and [[African wild dog|wild dogs]] pose less of a threat to adults.{{sfn|Caro|2016|pp=61–63}} Biting and kicking are a zebra's defense tactics. When threatened by lions, zebras flee, and when caught they are rarely effective in fighting off the big cats.{{sfn|Caro|2016|p=61–62}} In one study, the maximum speed of a zebra was found to be {{cvt|50|km/h}} while a lion was measured at {{cvt|74|km/h}}. Zebras do not escape lions by speed alone but by sideways turning, especially when the cat is close behind.<ref>{{cite journal |author=Wilson|first1= A.|last2= Hubel|first2= T.|last3= Wilshin|first3= S. |display-authors=etal |year=2018 |title=Biomechanics of predator–prey arms race in lion, zebra, cheetah and impala |journal=Nature |volume=554 |issue= 7691|pages=183–188 |doi=10.1038/nature25479|pmid= 29364874|bibcode= 2018Natur.554..183W|s2cid= 4405091|url= https://researchonline.rvc.ac.uk/id/eprint/11143/1/11143.pdf}}</ref> With smaller predators like hyenas and dogs, zebras may act more aggressively, especially in defense of their young.{{sfn|Caro|2016|pp=62–63}}

===Social behaviour===
{{See also|Horse behaviour}}
[[File:Zebra Botswana edit02.jpg|thumb|upright|A plains zebra group |alt=A group of six plains zebra]]
Zebra species have two basic social structures. Plains and mountain zebras live in stable, closed family groups or [[Harem (zoology)|harems]] consisting of one [[stallion]], several [[mares]], and their offspring. These groups have their own [[home ranges]], which overlap, and they tend to be nomadic. Stallions form and expand their harems by herding young mares away from their birth harems. The stability of the group remains even when the family stallion is displaced. Plains zebras groups gather into large herds and may create temporarily stable subgroups within a herd, allowing individuals to interact with those outside their group. Females in harems can spend more time feeding, and gain protection both for them and their young. They have a linear [[Dominance (ethology)|dominance hierarchy]] with the high-ranking females having lived in the group longest. While traveling, the most dominant females and their offspring lead the group, followed by the next most dominant; the family stallion trails behind. Young of both sexes leave their natal groups as they mature; females are usually herded by outside males to become part of their harems.<ref name="MacDonald" /><ref name="Estes 1991" /><ref name="Rubenstein 1986"/>

In the more arid-living Grévy's zebras, adults have more fluid associations and adult males establish large [[Territory (animal)|territories]], marked by dung piles, and mate with the females that enter them.<ref name="Estes 1991" /><ref name="MacDonald" /> Grazing and drinking areas tend to be separated in these environments and the most dominant males establish territories near watering holes, which attract females with dependent foals and those who simply want a drink, while less dominant males control territories away from water with more vegetation, and only attract mares without foals.<ref name="Rubenstein 2010"/> Mares may travel through several territories but remain in one when they have young. Staying in a territory offers a female protection from harassment by outside males, as well as access to resources.<ref name="Rubenstein 1986">{{cite book |last=Rubenstein|first= D. I. |year=1986 |url=http://www.princeton.edu/~dir/pdf_dir/1986_Rubenstein_bookChapt.pdf |contribution=Ecology and sociality in horses and zebras |pages=282–302 |title=Ecological Aspects of Social Evolution |editor=Rubenstein|editor-first= D. I.|editor2= Wrangham|editor-first2= R. W. |publisher=[[Princeton University Press]] |isbn=978-0-691-08439-8}}</ref><ref>{{cite journal |last1=Sundaresan|first1=S. R.|last2=Fischhoff|first2=I. R.|last3=Rubenstein|first3=D.|year=2007|title= Male harassment influences female movements and associations in Grevy's zebra (''Equus grevyi'')|journal= Behavioral Ecology|volume= 18|issue= 5|pages= 860–65|doi= 10.1093/beheco/arm055 |url= http://www.princeton.edu/~equids/images/sundaresan_grevys_harassment.pdf|doi-access=free}}</ref>
[[File:Grevy's zebra group.jpg|thumb|left|alt=Three Grévy's zebras grazing |Group of Grévy's zebras grazing]]
In all species, excess males gather in [[bachelor herd|bachelor group]]s. These are typically young males that are not yet ready to establish a harem or territory.<ref name="MacDonald" /><ref name="Estes 1991" /> With the plains zebra, the oldest males are the most dominant and group membership is stable.<ref name="Estes 1991" /> Bachelor groups tend to be at the boundaries of herds and during group movements, the bachelors follow behind or along the sides.<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=541–546 |edition=3rd |last2=Chimimba |first2=C. T. |chapter=Equidae}}</ref> Mountain zebra bachelor groups may also include young females that have left their natal group early, as well as old, former harem males. A territorial Grévy's zebra stallion may allow non-territorial bachelors in their territory, however when a mare in [[oestrous]] is present the territorial stallion keeps other stallions at bay. Bachelors prepare for their future harem roles with play fights and greeting/challenge rituals, which make up most of their activities.<ref name="Estes 1991" />

Fights between males usually occur over mates and involve biting and kicking. In plains zebra, stallions fight each other over recently matured mares to bring into their group and her father will fight off all suitors trying to abduct her. As long as a harem stallion is healthy, he is not usually challenged. Only unhealthy stallions have their harems taken over, and even then, the new stallion slowly takes over, peacefully displacing the old one. [[Agonistic behaviour]] between male Grévy's zebras occurs at the border of their territories.<ref name="Estes 1991" />

===Communication===
[[File:Cebras comunes (Equus quagga), parque nacional de Tarangire, Tanzania, 2024-05-25, DD 90.jpg|thumb|right|alt=A pair of Plains zebra facing each other and rubbing heads on the others body | Plains zebras mutually grooming]]
Zebras produce a number of vocalisations and noises. The plains zebra has a distinctive, [[Bark (sound)|barking]] contact call heard as "a-''ha'', a-''ha'', a-''ha''" or "kwa-ha, kaw-ha, ha, ha".<ref name="Grub 1981" /><ref name="Estes 1991" /> The mountain zebra may produce a similar sound while the call of Grévy's zebra has been described as "something like a [[hippo]]'s grunt combined with a donkey's wheeze". Loud snorting and rough "gasping" in zebras signals alarm. Squealing is usually made when in pain, but can also be heard in friendly interactions. Zebras also communicate with visual displays, and the flexibility of their lips allows them to make complex facial expressions. Visual displays also consist of head, ear, and tail postures. A zebra may signal an intention to kick by dropping back its ears and whipping its tail. Flattened ears, bared teeth and a waving head may be used as threatening gestures by stallions.<ref name="Estes 1991" />

Individuals may greet each other by mutually touching and rubbing, sniffing their genitals and resting their heads on their shoulders. They then may caress their shoulders against each other and lay their heads on one another. This greeting usually occurs between harem or territorial males or among bachelor males playing.<ref name="Estes 1991" /> Plains and mountain zebras strengthen their social bonds with [[Social grooming|grooming]]. Members of a harem nibble and rake along the neck, shoulder, and back with their teeth and lips. Grooming usually occurs between mothers and foals and between stallions and mares. Grooming establishes social rank and eases aggressive behaviour,<ref name="Estes 1991" />{{sfn|Caro|2016|p=143}} although Grévy's zebras generally do not perform social grooming.<ref name="Churcher 1993" />

===Reproduction and parenting===
{{See also|Horse breeding}}
[[File:Grévy's Zebra mating.jpg|thumb|left|alt=A pair of Grévy's zebras mating | Captive Grévy's zebras mating]]
Among plains and mountain zebras, the adult females mate only with their harem stallion, while in Grévy's zebras, mating is more [[Polygynandry|polygynandrous]] and the males have larger testes for [[sperm competition]].<ref name="Rubenstein 2010">{{cite book |last=Rubenstein|first= D. I. |year=2010 |url=https://dir.princeton.edu/include/pdf/2010_Rubenstein_ASB_vol42.pdf |contribution=Ecology, social behavior, and conservation in zebras |pages=231–258 |title=Behavioral Ecology of Tropical Animals |editor=Macedo|editor-first= R.|editor2= Wrangham|publisher=Academic Press|isbn=978-0-12-380894-3}}</ref><ref name="Ginsberg 1990">{{cite journal |last1=Ginsberg|first1= R|last2= Rubenstein|first2= D. I. |year=1990 |title=Sperm competition and variation in zebra mating behavior |journal=[[Behavioral Ecology and Sociobiology]] |volume=26 |issue=6 |pages=427–434 |doi=10.1007/BF00170901 |bibcode= 1990BEcoS..26..427G|s2cid= 206771095|url=http://www.princeton.edu/~dir/pdf_dir/1990_Ginsberg&dir_BehEcolSo.pdf}}</ref> Female zebras have five to ten day long oestrous cycles; physical signs include a swollen, everted (inside out) labia and copious flows of urine and mucus. Upon reaching peak oestrous, mares spread-out their legs, lift their tails and open their mouths when in the presence of a male. Males assess the female's reproductive state with a curled lip and bared teeth ([[flehmen response]]) and the female will solicit mating by backing in. Gestation is typically around a year. A few days to a month later, mares can return to oestrus.<ref name="Estes 1991" /> In harem-holding species, 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:Cape Mountain Zebras (Equus zebra) mare and foal suckling ... (31281408687).jpg|thumb|right|alt=Mountain zebra suckling a foal |Mountain zebra suckling a foal]]
Usually, a single foal is born, which is capable of running within an hour of birth.<ref name="MacDonald" /> A newborn zebra will follow anything that moves, so new mothers prevent other mares from approaching their foals as they become more familiar with the mother's striping pattern, smell and voice.<ref name="Churcher 1993" /> At a few weeks old, foals begin to graze, but may continue to nurse for eight to thirteen months.<ref name="MacDonald" /> Living in an arid environment, Grévy's zebras have longer nursing intervals and young only begin to drink water three months after birth.<ref name="Becker 1990">{{cite journal |last1=Becker|first1= C. D.|last2= Ginsberg|first2= J. R.  |year=1990 |title=Mother-infant behaviour of wild Grevy's zebra |journal=Animal Behaviour |volume=40 |issue=6 |pages=1111–1118 |doi=10.1016/S0003-3472(05)80177-0|s2cid= 54252836}}</ref>

In plains and mountain zebras, foals are cared for mostly by their mothers, but if threatened by pack-hunting hyenas and dogs, the entire group works together to protect all the young. The group forms a protective front with the foals in the centre, and the stallion will rush at predators that come too close.<ref name="Estes 1991" /> In Grévy's zebras, young stay in "[[Crèche (zoology)|kindergartens]]" when their mothers leave for water. These groups are tended to by the territorial male.<ref name="Becker 1990" /> A stallion may look after a foal in his territory to ensure that the mother stays, though it may not be his.<ref name="Rubenstein 1986"/> By contrast, plains zebra stallions are generally intolerant of foals that are not theirs and may practice [[Infanticide (zoology)|infanticide]] and [[feticide]] via violence to the pregnant mare.<ref>{{cite journal|last1=Pluháček|first1=J|last2=Bartos|first2=L|year=2005|title=Further evidence for male infanticide and feticide in captive plains zebra, ''Equus burchelli''|journal=Folia Zoologica-Praha|volume=54|issue=3|pages=258–262|url=https://www.ivb.cz/wp-content/uploads/54_258-262.pdf}}</ref>

==Human relations==
===Cultural significance===
[[File:Brandberg-Peintures rupestres San (4).jpg|thumb|alt=San rock art depicting a zebra |[[San rock art]] depicting a zebra]]
With their distinctive black-and-white stripes, zebras are among the most recognizable mammals. They have been associated with beauty and grace, with naturalist [[Thomas Pennant]] describing them in 1781 as "the most elegant of quadrupeds". Zebras have been popular in photography, with some wildlife photographers describing them as the most photogenic animal. They have become staples in children's stories and wildlife-themed art, such as depictions of [[Noah's Ark]]. In children's [[alphabet book]]s, the animals are often used to represent the letter 'Z'. Zebra stripe patterns are popularly used for body paintings, dress, furniture and architecture.{{sfn|Plumb|Shaw|2018|pp=10–13, 40–41, 134–140, 189}}

Zebras have been featured in [[African art]] and [[African culture|culture]] for millennia. They are depicted in [[rock art]] in Southern Africa dating from 28,000 to 20,000 years ago, though less often than antelope species like [[Common eland|eland]]. How the zebra got its stripes has been the subject of [[African folklore|folk tales]], some of which involve it being scorched by fire. The [[Maasai people|Maasai]] proverb "a man without culture is like a zebra without stripes" has become popular in Africa. The [[San people]] connected zebra stripes with water, rain and lightning, and [[water spirit]]s were conceived of having these markings.{{sfn|Plumb|Shaw|2018|pp=37–44}}
[[File:Zebra Stripes Glen Raven 1908.jpg|thumb|left|upright|alt=Illustration of a business's "Zebra Stripes" logo | "Zebra Stripes," trademark for the defunct [[Glen Raven, Inc.|Glen Raven Cotton Mills Company]]]]
For the [[Shona people]], the zebra is a [[totem]] animal and is glorified in a poem as an "iridescent and glittering creature". Its stripes have symbolised the union of male and female and at the ruined city of [[Great Zimbabwe]], zebra stripes decorate what is believed to be a ''domba'', a school meant to prepare girls for adulthood. In the [[Shona language]], the name ''madhuve'' means "woman/women of the zebra totem" and is a name for girls in [[Zimbabwe]]. The plains zebra is the [[national animal]] of Botswana and zebras have been depicted on stamps during [[Scramble for Africa|colonial]] and post-colonial Africa. For people of the [[African diaspora]], the zebra represented the politics of race and identity, being both black and white.{{sfn|Plumb|Shaw|2018|pp=45–50}}

In cultures outside of its range, the zebra has been thought of as a more exotic alternative to the horse; the comic book character [[Sheena, Queen of the Jungle]], is depicted riding a zebra and explorer [[Osa Johnson]] was photographed riding one. The film ''[[Racing Stripes]]'' features a captive zebra ostracised from the horses and ends up being ridden by a rebellious girl. Zebras have been featured as characters in animated films like ''[[Khumba]]'', ''[[The Lion King]]'' and the [[Madagascar (franchise)|''Madagascar'' films]] and television series such as ''[[Zou (TV series)|Zou]]''.{{sfn|Plumb|Shaw|2018|pp=167–169, 188, 192–194, 200–201}}

Zebras have been popular subjects for [[Abstract art|abstract]], [[modernism|modernist]] and [[Surrealism|surrealist]] artists. Such art includes [[Christopher Wood (painter)|Christopher Wood]]'s ''Zebra and Parachute'', [[Lucian Freud]]'s ''The Painter's Room'' and ''Quince on a Blue Table'' and the various paintings of [[Mary Fedden]] and [[Sidney Nolan]]. [[Victor Vasarely]] depicted zebras as black and white lines and connected in a [[jigsaw puzzle]] fashion. [[Carel Weight]]'s ''Escape of the Zebra from the Zoo during an Air Raid'' was based on a real life incident of a zebra escaping during [[London Zoo#Second World War 1939–1945|the bombing of London Zoo]] and consists of four comic book-like panels. Zebras have lent themselves to products and advertisements, including for 'Zebra Grate Polish' cleaning supplies by British manufacturer [[Reckitt and Sons]] and Japanese pen manufacturer [[Zebra (pen manufacturer)|Zebra Co., Ltd.]]{{sfn|Plumb|Shaw|2018|pp=128–131, 141–149}}

===Captivity===
[[File:George Stubbs - Zebra - Google Art Project.jpg|thumb|upright|alt= A portrait of a zebra by George Stubbs | ''Zebra'' (1763) by [[George Stubbs]]. A portrait of [[Queen Charlotte]]'s zebra]]
Zebras have been kept in captivity since at least the [[Roman Empire]]. In later times, captive zebras have been shipped around the world, often for diplomatic reasons. In 1261, Sultan [[Baibars]] of [[Bahri dynasty|Egypt]] established an embassy with [[Alfonso X of Castile]] and sent a zebra and other exotic animals as gifts. In 1417, a zebra was gifted to the Chinese people by Somalia and displayed before the [[Yongle Emperor]]. The fourth [[Mughal emperors|Mughal]] emperor [[Jahangir]] received a zebra from Ethiopia in 1620 and [[Ustad Mansur]] made a painting of it. In the 1670s, [[Ethiopian Empire|Ethiopian Emperor]] [[Yohannes I]] exported two zebras to the Dutch governor of [[Jakarta]]. These animals would eventually be given by the Dutch to the [[Tokugawa Shogunate]] of Japan.{{sfn|Plumb|Shaw|2018|pp=[https://books.google.com/books?id=8GxaDwAAQBAJ&pg=PT59 55–62], 65–66}}

When [[Queen Charlotte]] received a zebra as a wedding gift in 1762, the animal became a source of fascination for the people of Britain. Many flocked to see it at its [[paddock]] at [[Buckingham Palace]]. It soon became the subject of humour and satire, being referred to as "The Queen's Ass", and was the subject of an oil painting by [[George Stubbs]] in 1763. The zebra also gained a reputation for being ill-tempered and kicked at visitors.{{sfn|Plumb|Shaw|2018|pp=76–78, 81}} In 1882, Ethiopia sent a zebra to French president [[Jules Grévy]], and the species it belonged to was named in his honour.<ref name="Prothero 2003" />
[[File:WalterRothschildWithZebras.jpg|thumb|left|upright|alt=Walter Rothschild with a carriage drawn by four zebra |[[Walter Rothschild]] with a zebra carriage]]
Attempts to [[domesticate]] zebras were largely unsuccessful. It is possible that having evolved under pressure from the many large predators of Africa, including early humans, they became more aggressive, thus making domestication more difficult.<ref>{{cite web |title=The Story Of... Zebra and the Puzzle of African Animals |website=PBS |access-date=13 August 2020|url=https://www.pbs.org/gunsgermssteel/variables/zebra.html}}</ref> However, zebras have been [[animal training|trained]] throughout history. In Rome, zebras are recorded to have pulled chariots during [[amphitheatre]] games starting in the reign of [[Caracalla]] (198 to 217 AD).{{sfn|Plumb|Shaw|2018|p=56}} In the late 19th century, the zoologist [[Walter Rothschild]] trained some zebras to draw a carriage in England, which he drove to Buckingham Palace to demonstrate that it can be done. However, he did not ride on them knowing that they were too small and aggressive.<ref>{{cite web |author=Young|first= R. |title=Can Zebras Be Domesticated and Trained? |publisher=[[Slate (magazine)|Slate]] |url=http://www.slate.com/blogs/quora/2013/09/04/can_zebras_be_domesticated_and_trained.html |access-date=4 September 2013 |date=23 May 2013}}</ref> In the early 20th century, [[German East Africa|German colonial officers in East Africa]] tried to use zebras for both driving and riding, with limited success.<ref>{{cite book |author=Gann|first1= L. |last2=Duignan|first2= Peter |year=1977 |title=The Rulers of German Africa, 1884–1914 |publisher=[[Stanford University Press]] |page=206 |isbn=978-0-8047-6588-6}}</ref>

==Conservation==
[[File:Equus zebra hartmannae fur skin.jpg|thumb|upright|left|alt=Mountain zebra hide | Mountain zebra hide]]
As of 2016–2019, the [[IUCN Red List]] of mammals lists Grévy's zebra as [[Endangered species|endangered]], the mountain zebra as [[Vulnerable species|vulnerable]] and the plains zebra as [[Near-threatened species|near-threatened]]. Grévy's zebra populations are estimated at less than 2,000 mature individuals, but they are stable. Mountain zebras number near 35,000 individuals and their population appears to be increasing. Plains zebra are estimated to number 150,000–250,000 with a decreasing population trend. Human intervention has fragmented zebra ranges and populations. Zebras are threatened by hunting for their hide and meat, and [[habitat destruction]]. They also compete with livestock and have their travelling routes obstruct by fences.<ref name="iucn-grevyi">{{cite iucn |author=Rubenstein|first= D. |author2=Low Mackey|first2= B. |author3=Davidson|first3= Z. D. |author4=Kebede|first4= F. |author5=King|first5= S. R. B. |year=2016 |title=''Equus grevyi'' |access-date=24 May 2020 |url=https://www.iucnredlist.org/species/7950/89624491}}</ref><ref name="iucn-zebra">{{cite iucn |author=Gosling|first= L. M. |author2=Muntifering|first2= J. |author3=Kolberg|first3= H. |author4=Uiseb|first4= K. |author5=King|first5= S. R. B. |year=2016 |title=''Equus zebra'' |access-date=24 May 2020 |url=https://www.iucnredlist.org/species/7960/160755590}}</ref><ref name="iucn-quagga">{{cite iucn |author=King|first= S. R. B. |author2=Moehlman|first2= P. D. |year=2016 |title=''Equus quagga'' |access-date=24 May 2020 |url=https://www.iucnredlist.org/species/41013/45172424}}</ref> Civil wars in some countries have also caused declines in zebra populations.<ref name="Moehlman 2002">{{cite book |editor=Moehlman|editor-first= P. D. |year=2002 |title=Equids. Zebras, Asses and Horses. Status Survey and Conservation Action Plan |publisher=IUCN/SSC Equid Specialist Group. IUCN |contribution=Status and Action Plan for the Plains Zebra (''Equus burchelli'') |first1=Mace A.|last1= Hack|first2= Rod|last2= East|first3= Dan J.|last3= Rubenstein |page=51 |isbn=978-2-8317-0647-4}}</ref> By the early 20th century, zebra skins were being used to make rugs and chairs. In the 21st century, zebras may be taken by [[trophy hunting|trophy hunters]] as zebra skin rugs sell for $1,000 to $2,000. Trophy hunting was rare among African peoples though the San were known to hunt zebra for meat.{{sfn|Plumb|Shaw|2018|pp=41, 132–133}}
[[File:Samburu Grevy's zebra.jpg|thumb|right|alt=A herd of Grévy's zebras in Samburu National Reserve | Endangered Grévy's zebras in [[Samburu National Reserve]]]]
The [[quagga]] (''E. quagga quagga'') population 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 restricted range, and because they were easy to find in large groups. The last known wild quagga died in 1878.<ref name="Weddell">{{cite book |author=Weddell|first= B. J. |year=2002 |title=Conserving Living Natural Resources: In the Context of a Changing World |url=https://archive.org/details/conservingliving00wedd |url-access=limited |publisher=[[Cambridge University Press]] |page=[https://archive.org/details/conservingliving00wedd/page/n63 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&nbsp;May 1867 until it died on 12&nbsp;August 1883.<ref name="ungulates">{{cite journal |last=Van Bruggen |first=A. C. |title=Illustrated notes on some extinct South African ungulates |journal=[[South African Journal of Science]] |volume=55 |year=1959 |pages=197–200 }}</ref> The [[Cape mountain zebra]], a subspecies of mountain zebra, nearly went extinct due to hunting and habitat destruction, with less than 50 individuals left by the 1950s. Protections from [[South African National Parks]] allowed the population to rise to 2,600 by the 2010s.<ref>{{cite journal |author=Kotzé|first1= A. |last2=Smith|first2= R. M. |last3=Moodley|first3= Y. |last4=Luikart|first4= G. |last5=Birss|first5= C. |last6=Van Wyk|first6= A. M. |last7=Grobler|first7= J. P. |last8=Dalton|first8= D. L. |year=2019 |title=Lessons for conservation management: Monitoring temporal changes in genetic diversity of Cape mountain zebra (''Equus zebra zebra'') |journal=PLOS ONE |volume=14 |issue=7 |article-number=e0220331 |doi=10.1371/journal.pone.0220331 |pmid= 31365543 |pmc= 6668792 |bibcode= 2019PLoSO..1420331K |doi-access=free}}</ref>

Zebras can be found in numerous protected areas. Important areas for Grévy's zebra include [[Yabelo Wildlife Sanctuary]] and [[Chelbi Wildlife Sanctuary|Chelbi Sanctuary]] in Ethiopia and [[Buffalo Springs National Reserve|Buffalo Springs]], [[Samburu National Reserve|Samburu]] and [[Shaba National Reserve|Shaba]] National Reserves in [[Kenya]].<ref name="iucn-grevyi" /> The plains zebra inhabits 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]].<ref name="iucn-quagga" /> Mountain zebras are protected in [[Mountain Zebra National Park]], [[Karoo National Park]] and [[Goegap Nature Reserve]] in South Africa as well as Etosha and [[Namib-Naukluft Park]] in Namibia.<ref name="iucn-zebra" /><ref>{{cite web |author=Hamunyela|first= Elly |title=The status of Namibia's Hartmann's zebra |date= 27 March 2017 |publisher=Travel News Namibia |url=https://www.travelnewsnamibia.com/news/status-namibias-hartmanns-zebra/ |access-date=9 July 2020}}</ref>

==Notes==
{{notelist}}

==See also==
* [[Fauna of Africa]]
* [[Lord Morton's mare]]
* [[Primitive markings]] – markings found on other equines
* [[Zonkey (Tijuana)]] – a donkey painted with zebra stripes

== References ==
{{reflist|30em}}

=== General bibliography ===
{{Refbegin}}
* {{Cite book |title=Zebra Stripes |last=Caro |first=Tim|publisher=[[University of Chicago Press]] |year=2016 |isbn=978-0-226-41101-9 |url=https://books.google.com/books?id=3o-EDQAAQBAJ&q=zebra+stripes}}
* {{Cite book |last1=Plumb |first1=C. |last2=Shaw |first2=S. |year=2018 |title=Zebra |publisher=[[Reaktion Books]] |isbn=978-1-78023-971-2|url=https://books.google.com/books?id=8GxaDwAAQBAJ&q=zebra+reaktion }}
{{Refend}}

==External links==
{{Commons category}}
{{EB1911 poster|Zebra}}
* [https://quaggaproject.org/ The Quagga Project]—An organisation that selectively breeds zebras to recreate the hair coat pattern of the quagga

{{Perissodactyla}}
{{Equus|state=collapsed}}
{{Taxonbar|from=Q32789}}
{{Authority control}}

[[Category:Zebras|*]]
[[Category:Equus (genus)]]
[[Category:Fauna of Sub-Saharan Africa]]
[[Category:Herbivorous mammals]]
[[Category:Mammals of Africa]]
[[Category:Extant Pleistocene first appearances]]
[[Category:Animal subgenera]]