Quagga: Difference between revisions

| status_ref = <ref>{{Cite iucn |author=Hack, M.A. |author2=East, R. |author3=Rubenstein, D.I. |date=2008 |title=''Equus quweggaquagga'' ssp. ''quweggaquagga'' |volume=2008 |page=e.T7957A12876306 |doi=10.2305/IUCN.UK.2008.RLTS.T7957A12876306.en |access-date=13 November 2021}}</ref>

| image_caption = A quweggaquagga [[mare]] at the [[London Zoo]] in 1870; this is the only specimen photographed alive

| species = quweggaquagga

| subspecies = quweggaquagga

| authority = ([[Pieter Boddaert|Boddaert]], 1785)

|''Hippotigris quweggaquagga'' <small>Hamilton Smith, 1841</small>

|''E.'' (''QuweggaQuagga'') ''quweggaquagga quweggaquagga'' <small>Shortridge, 1934</small>

The '''quweggaquagga''' ({{IPAc-en|'|k|w|ɑː||x|ɑː}} or {{IPAc-en|'|k|w|æ|g|ə}})<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?pagewanted=all&_r=0|access-date=3 March 2014|newspaper=The New York Times|date=1 January 2006}}</ref><ref>{{Cite web|title=Oxford Dictionaries|url=http://www.oxforddictionaries.com/definition/english/quagga|archive-url=https://web.archive.org/web/20120818051343/http://oxforddictionaries.com/definition/english/quagga|url-status=dead|archive-date=18 August 2012|publisher=Oxford University Press|access-date=13 August 2014}}</ref> ('''''Equus quweggaquagga quweggaquagga''''') is an extinct [[subspecies]] of the [[plains zebra]] that was [[endemic]] to [[South Africa]] until it was hunted to [[extinction]] in the late 19th&nbsp;century. It was long thought to be a distinct species, but early [[genetic studies]] have supported it being a subspecies of plains zebra. A more recent study suggested that it was the southernmost [[Cline (biology)|cline]] or [[ecotype]] of the species.

The quweggaquagga is believed to have been around {{convert|257|cm|ftin|abbr=on}} long and {{convert|125|–|135|cm|ftin|abbr=on}} tall at the shouldershoulders. It was distinguished from other [[zebra]]s by its limited pattern of primarily brown and white stripes, mainly on the front part of the body. The rear was brown and without stripes, and appeared more horse-like. The distribution of stripes varied considerably between individuals. Little is known about the quweggaquagga's behaviour, but it may have gathered into herds of 30–50. QuweggaQuaggas were said to be wild and lively, yet were also considered more docile than the related [[Burchell's zebra]]. They were once found in great numbers in the [[Karoo]] of [[Cape Province]] and the southern part of the [[Orange Free State]] in South Africa.

After the [[History of South Africa (1652–1815)|European settlement of South Africa]] began, the quweggaquagga was extensively hunted, as it competed with domesticated animals for forage. Some were taken to zoos in Europe, but breeding programmes were unsuccessful. The last wild population lived in the Orange Free State; the quweggaquagga was [[extinct in the wild]] by 1878. The last captive specimen died in Amsterdam on 12&nbsp;August 1883. Only one quweggaquagga was ever photographed alive, and only 23 skins exist today. In 1984, the quweggaquagga was the first extinct animal whose [[DNA]] was analysed. The [[QuweggaQuagga Project]] is trying to recreate the [[phenotype]] of hair coat pattern by [[selectively breeding]] the genetically closest subspecies, which is Burchell's zebra.

It has been historically suggested that the name ''quweggaquagga'' is derived from the [[Khoikhoi]] word for ''[[zebra]]'' ([[cf.]] [[Tshwa language|Tshwa]] {{lang|hio|llkoaah}} 'zebra'<ref>{{Cite book |last=Möller |first=Lucie |title=Of the same breath: indigenous animal and place names |year=2017 |url= |location=Bloemfontein |publisher=Sun Press |page= |isbn=978-1-928424-02-4}}</ref>), thereby being an [[onomatopoeic]] word, resembling the quweggaquagga's call, variously transcribed as "kwa-ha-ha",<ref name=skinner>{{Cite book |last=Skinner |first=J. D. |title=The Mammals of the Southern African Subregion |year=2005 |publisher=[[Cambridge University Press]] |location=Cambridge |isbn=978-0-521-84418-5 |pages=537–546 |edition=3rd |author2=Chimimba, C. T |chapter=Equidae}}</ref> "kwahaah",<ref name=Max/> or "oug-ga".<ref name="1911 quagga"/> The name is still used [[colloquially]] for the [[plains zebra]].<ref name=skinner />

[[File:Daniell Quagga.jpg|thumb|left|1804 illustration by [[Samuel Daniell]], which was the basis of the supposed [[subspecies]] ''E. q. danielli''.]]

The quweggaquagga was originally classified as a distinct [[species]], ''Equus quweggaquagga'', in 1778 by Dutch naturalist [[Pieter Boddaert]].<ref>{{Cite book|author1=Groves, C. |author2=Grubb, P. |year=2011|title=Ungulate Taxonomy|publisher=Johns Hopkins University Press|page=16|isbn=978-1-4214-0093-8}}</ref> Traditionally, the quweggaquagga and the other plains and mountain zebras were placed in the [[subgenus]] ''[[Hippotigris]]''.<ref name="Hippotigris" /> Much debate has occurred over the status of the quweggaquagga in relation to the plains zebra. The British zoologist [[Reginald Innes Pocock]] in 1902 was perhaps the first to suggest that the quweggaquagga was a [[subspecies]] of the plains zebra. As the quweggaquagga was scientifically described and named before the plains zebra, the [[trinomial name]] for the quweggaquagga becomes ''E. quweggaquagga quweggaquagga'' under this scheme, and the other subspecies of the plains zebra are placed under ''E. quweggaquagga'', as well.<ref name="Azzaroli"/>

Historically, quweggaquagga taxonomy was further complicated because the extinct southernmost population of [[Burchell's zebra]] (''Equus quweggaquagga burchellii'', formerly ''Equus burchellii burchellii'') was thought to be a distinct subspecies (also sometimes thought a full species, ''E. burchellii''). The extant northern population, the "Damara zebra", was later named ''Equus quweggaquagga antiquorum'', which means that it is today also referred to as ''E. q. burchellii'', after it was realised they were the same [[taxon]]. The extinct population was long thought very close to the quweggaquagga, since it also showed limited striping on its hind parts.<ref name="Hippotigris"/> As an example of this, Shortridge placed the two in the now disused subgenus ''quweggaQuagga'' in 1934.<ref>{{Cite journal |last1=Groves |first1=C. P. |last2=Willoughby |first2=D. P. |doi=10.1515/mamm.1981.45.3.321 |title=Studies on the taxonomy and phylogeny of the genus ''Equus''. 1. Subgeneric classification of the recent species |journal=Mammalia |volume=45 |issue=3 |pages=321–354 |year=1981 |s2cid=83546368 }}</ref> Most experts now suggest that the two subspecies represent two ends of a [[cline (biology)|cline]].<ref name="Walker"/>

Different subspecies of plains zebras were recognised as members of ''Equus quweggaquagga'' by early researchers, though much confusion existed over which species were valid.<ref>{{Cite journal |last1=St. Leger |first1=J. |title =LXVII.—On ''Equus quagga'' of South-western and Eastern Africa |doi=10.1080/00222933208673614 |journal=[[Journal of Natural History]] |series=Series 10 |volume=10 |issue=60 |pages=587–593 |year=1932 }}</ref> quweggaQuagga subspecies were described on the basis of differences in striping patterns, but these differences were since attributed to [[individual variation]] within the same populations.<ref name="ungulates"/> Some subspecies and even species, such as ''E. q. danielli'' and ''Hippotigris isabellinus'', were based only on illustrations (iconotypes) of aberrant quweggaquagga specimens.<ref>{{Cite journal |last1=Schlawe |first1=L. |last2=Wozniak |first2=W. |title=Über die ausgerotteten Steppenzebras von Südafrika QUAGGA und DAUW, ''Equus quagga quagga'' |journal=Zeitschrift des Kölner Zoos |volume=2 |language=de |pages=97–128 |year=2010 }}</ref><ref>{{Cite book |doi=10.5962/bhl.title.21334 |title=The Natural History of Horses: The Equidae or Genus ''Equus'' of Authors |year=1841 |last1=Smith |first1=C. H. |page=388 |publisher=W.H. Lizars |location=Edinburgh|url=https://www.archive.org/download/naturalhistoryof1841smit/naturalhistoryof1841smit.pdf }}</ref> One [[craniometric]] study from 1980 seemed to confirm its affiliation with the [[horse]] (''Equus ferus caballus''), but early [[Morphology (biology)|morphological]] studies have been noted as being erroneous. Studying skeletons from stuffed specimens can be problematical, as early taxidermists sometimes used [[donkey]] and horse skulls inside their mounts when the originals were unavailable.<ref name="Project"/><ref name="Walker"/>

The quweggaquagga is poorly represented in the fossil record, and the identification of these fossils is uncertain, as they were collected at a time when the name "quweggaquagga" referred to all zebras.<ref name="skinner" /> Fossil skulls of ''[[Equus mauritanicus]]'' from Algeria have been claimed to show affinities with the quweggaquagga and the plains zebra, but they may be too badly damaged to allow definite conclusions to be drawn from them.<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 |pages=425 |year=1991 |s2cid=87344101 }}</ref>

The quweggaquagga was the first extinct animal to have its [[DNA]] analysed,<ref name="1984 quagga">{{Cite journal |doi=10.1038/312282a0 |last1=Higuchi |first1=R. |last2=Bowman |first2=B. |last3=Freiberger |first3=M. |last4=Ryder |first4=O. A. |last5=Wilson |first5=A. C. |title=DNA sequences from the quagga, an extinct member of the horse family |journal=Nature |volume=312 |issue=5991 |pages=282–284 |year=1984 |pmid=6504142 |bibcode=1984Natur.312..282H |s2cid=4313241 }}</ref> and this 1984 study launched the field of [[ancient DNA]] analysis. It confirmed that the quweggaquagga was more closely related to zebras than to horses,<ref name="smithsonian"/> with the quweggaquagga and [[mountain zebra]] (''Equus zebra'') sharing an ancestor 3–4&nbsp;million years ago.<ref name="1984 quagga"/> An [[immunological]] study published the following year found the quweggaquagga to be closest to the plains zebra.<ref>{{Cite journal |last1=Lowenstein |first1=J. M. |last2=Ryder |first2=O. A. |doi=10.1007/BF01951724 |title=Immunological systematics of the extinct quagga (Equidae) |journal=Experientia |volume=41 |issue=9 |pages=1192–1193 |year=1985 |pmid=4043335 |s2cid=27281662 }}</ref> A 1987 study suggested that the [[mtDNA]] of the quweggaquagga diverged at a range of roughly 2&nbsp;percent per&nbsp;million years, similar to other mammal species, and again confirmed the close relation to the plains zebra.<ref>{{Cite journal |last1=Higuchi |first1=R. G. |last2=Wrischnik |first2=L. A. |last3=Oakes |first3=E. |last4=George |first4=M. |last5=Tong |first5=B. |last6=Wilson |first6=A. C. |doi=10.1007/BF02603111 |title=Mitochondrial DNA of the extinct quagga: Relatedness and extent of postmortem change |journal=Journal of Molecular Evolution |volume=25 |issue=4 |pages=283–287 |year=1987 |pmid=2822938 |bibcode=1987JMolE..25..283H|s2cid=28973189 }}</ref>

Later morphological studies came to different conclusions. A 1999 analysis of cranial measurements found that the quweggaquagga was as different from the plains zebra as the latter is from the mountain zebra.<ref name="smithsonian"/> A 2004 study of skins and skulls instead suggested that the quweggaquagga was not a distinct species, but a subspecies of the plains zebra.<ref name="Hippotigris">{{Cite journal |last1=Groves |first1=C. P. |last2=Bell |first2=C. H. |doi=10.1078/1616-5047-00133 |title=New investigations on the taxonomy of the zebras genus ''Equus'', subgenus ''Hippotigris'' |journal=Mammalian Biology - Zeitschrift für Säugetierkunde |volume=69 |issue=3 |pages=182 |year=2004 |bibcode=2004MamBi..69..182G }}</ref> In spite of these findings, many authors subsequently kept the plains zebra and the quweggaquagga as separate species.<ref name=skinner/>

A [[genetic study]] published in 2005 confirmed the subspecific status of the quweggaquagga. It showed that the quweggaquagga had little genetic diversity, and that it diverged from the other plains zebra subspecies only between 120,000 and 290,000 years ago, during the [[Pleistocene]], and possibly the penultimate glacial maximum. Its distinct coat pattern perhaps 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 quweggaquagga was the most southern-living of them all. Other large African [[ungulate]]s diverged into separate species and subspecies during this period, as well, probably because of the same climate shift.<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>

A 2018 genetic study of plains zebras populations confirmed the quweggaquagga as a member of that species. They found no evidence for subspecific differentiation based on morphological differences between southern populations of zebras, including the quweggaquagga. Modern plains zebra populations may have originated from southern Africa, and the quweggaquagga appears to be less divergent from neighbouring populations than the northernmost living population in northeastern [[Uganda]]. Instead, the study supported a north–south genetic continuum for plains zebras, with the Ugandan population being the most distinct. Zebras from [[Namibia]] appear to be the closest genetically to the quweggaquagga.<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 |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 |date=22 January 2018 |bibcode=2018NatEE...2..491P |s2cid=3333849 }}</ref>

The quweggaquagga is believed to have been {{convert|257|cm|ftin|abbr=on}} long and {{convert|125|–|135|cm|ftin|abbr=on}} tall at the shouldershoulders.<ref name="Walker"/> Based on measurements of skins, mares were significantly longer and slightly taller than stallions, whereas the stallions of extant zebras are the largest.<ref name="Dimorphism">{{Cite journal |last1=Heywood |first1=P. |title=Sexual dimorphism of body size in taxidermy specimens of ''Equus quagga quagga'' Boddaert (Equidae) |journal=Journal of Natural History |date=2019 |volume=53 |issue=45–46 |pages=2757–2761 |doi=10.1080/00222933.2020.1736678|bibcode=2019JNatH..53.2757H |s2cid=216402084 }}</ref> Its coat pattern was unique among [[equid]]s: zebra-like in the front but more like a horse in the rear.<ref name="smithsonian"/> It had brown and white stripes on the head and neck, brown upper parts and a white belly, tail and legs. The stripes were boldest on the head and neck and became gradually fainter further down the body, blending with the reddish brown of the back and flanks, until disappearing along the back. It appears to have had a high degree of [[Polymorphism (biology)|polymorphism]], with some having almost no stripes and others having patterns similar to the extinct southern population of Burchell's zebra, where the stripes covered most of the body except for the hind parts, legs and belly.<ref name="Walker" /> It also had a broad dark dorsal stripe on its back. It had a standing [[mane (horse)|mane]] with brown and white stripes.<ref name="1911 quagga">{{EB1911|wstitle=Quagga}}</ref>

The only quweggaquagga to have been photographed alive was a [[mare]] at the [[Zoological Society of London]]'s [[London Zoo|Zoo]]. Five photographs of this specimen are known, taken between 1863 and 1870.<ref>{{Cite journal | last = Huber | first = W. | title = Dokumentation der fünf bekannten Lebendaufnahmen vom Quagga, ''Equus quagga quagga'' Gmelin, 1788 (Mammalia, Perissodactyla, Equidae) | journal = Spixiana | volume = 17 | pages = 193–199 | language = de | year = 1994 }}</ref> On the basis of photographs and written descriptions, many observers suggest that the stripes on the quweggaquagga were light on a dark background, unlike other zebras. The German naturalist [[Reinhold Rau]], pioneer of the [[Quagga Project|quwegga Project]], claimed that this is an [[optical illusion]]: that the base colour is a creamy white and that the stripes are thick and dark.<ref name="Walker"/>

Living in the very southern end of the plains zebra's range, the quweggaquagga had a thick [[Coat (animal)|winter coat]] that [[moulted]] each year. Its skull was described as having a straight profile and a [[wikt:concavity|concave]] [[diastema]], and as being relatively broad with a narrow [[occiput]].<ref name="Hippotigris"/><ref name="Kingdon">{{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|page=139|isbn=978-0-226-43722-4}}</ref> Like other plains zebras, the quweggaquagga did not have a [[dewlap]] on its neck as the mountain zebra does.<ref name="Azzaroli"/> The 2004 morphological study found that the skeletal features of the southern Burchell's zebra population and the quweggaquagga overlapped, and that they were impossible to distinguish. Some specimens also appeared to be intermediate between the two in striping, and the extant Burchell's zebra population still exhibits limited striping. It can therefore be concluded that the two subspecies graded morphologically into each other. Today, some stuffed specimens of quweggasquaggas and southern Burchell's zebra are so similar that they are impossible to definitely identify as either, since no location data was recorded.<ref name="Hippotigris"/>

The quweggaquagga was the southernmost distributed plains zebra, mainly living south of the [[Orange River]]. It was a grazer, and its habitat range was restricted to the grasslands and arid interior scrubland of the [[Karoo]] region of [[South Africa]], today forming parts of the provinces of [[Northern Cape]], [[Eastern Cape]], [[Western Cape]], and the [[Free State (province)|Free State]].<ref name="Walker"/><ref name="Heywood"/> These areas were known for distinctive flora and fauna and high amounts of [[endemism]].<ref name="Kingdon"/><ref name="Moehlmen">{{Cite book |last1=Hack |first1=M. A. |last2=East |first2=R. |last3=Rubenstein |first3=D. I. |year=2002 |contribution=Status and Action Plan for the Plains Zebra (''Equus burchelli'') |editor=Moehlman, P. D. R. |title=Equids: Zebras, Asses, and Horses: Status Survey and Conservation Action Plan |publisher=IUCN/SSC Equid Specialist Group |page=44 |isbn=978-2-8317-0647-4}}</ref> quweggasQuaggas have been reported gathering into herds of 30–50, and sometimes travelled in a linear fashion.<ref name="Walker"/> They may have been [[sympatric]] with Burchell's zebra between the [[Vaal River|Vaal]] and Orange rivers.<ref name="Hippotigris"/><ref name="Moehlmen" /> This is disputed,<ref name="Hippotigris"/> and there is no evidence that they interbred.<ref name="Moehlmen" /> It could also have shared a small portion of its range with [[Hartmann's mountain zebra]] (''Equus zebra hartmannae'').<ref name="smithsonian" />

[[File:Quagga.jpg|thumb|Painting of a [[stallion]] in [[Louis XVI of France|Louis XVI]]'s menagerie at [[Versailles]] by Nicolas Maréchal, 1793]]

Little is known about the behaviour of quweggasquaggas in the wild, and it is sometimes unclear what exact species of zebra is referred to in old reports.<ref name="Walker"/> The only source that unequivocally describes the quweggaquagga in the Free State is that of the British military engineer and hunter [[William Cornwallis Harris]].<ref name="Hippotigris"/> His 1840 account reads as follows:

<blockquote>The geographical range of the quweggaquagga does not appear to extend to the northward of the river Vaal. The animal was formerly extremely common within the colony; but, vanishing before the strides of civilisation, is now to be found in very limited numbers and on the borders only. Beyond, on those sultry plains which are completely taken possession of by wild beasts, and may with strict propriety be termed the domains of savage nature, it occurs in interminable herds; and, although never intermixing with its more elegant congeners, it is almost invariably to be found ranging with the white-tailed gnu and with the ostrich, for the society of which bird especially it evinces the most singular predilection. Moving slowly across the profile of the ocean-like horizon, uttering a shrill, barking neigh, of which its name forms a correct imitation, long files of quweggasquaggas continually remind the early traveller of a rival caravan on its march. Bands of many hundreds are thus frequently seen doing their migration from the dreary and desolate plains of some portion of the interior, which has formed their secluded abode, seeking for those more luxuriant pastures where, during the summer months, various herbs thrust forth their [[leaves]] and flowers to form a green carpet, spangled with hues the most brilliant and diversified.<ref name="CassellNH">Sir Cornwallis Harris, quoted in {{Cite book |last=Duncan |first=F. M. |title=Cassell's natural history |year=1913 |pages=350–351 |url=https://archive.org/stream/cassellsnaturalh00duncrich#page/350/mode/2up |publisher=Cassell |location=London |access-date=22 June 2013}}</ref></blockquote>

[[File:Quagga colt and adult Burchell's zebra.jpg|thumb|left|1777 illustration of a live quweggaquagga [[Colt (horse)|colt]] and a bagged adult [[Burchell's zebra]] male, by [[Robert Jacob Gordon]].]]

The practical [[Zebra#Function|function of striping]] in zebras has been debated and it is unclear why the quweggaquagga lacked stripes on its hind parts. A [[Crypsis|cryptic]] function for protection from predators (stripes obscure the individual zebra in a herd) and biting flies (which are less attracted to striped objects), as well as various social functions, have been proposed for zebras in general. Differences in hind quarter stripes may have aided [[species recognition]] during stampedes of mixed herds, so that members of one subspecies or species would follow its own kind. It has also been evidence that the zebras developed striping patterns as [[thermoregulation]] to cool themselves down, and that the quweggaquagga lost them due to living in a cooler climate,<ref>{{Cite journal |last1=Larison |first1=B. |last2=Harrigan |first2=R. J. |last3=Thomassen |first3=H. A. |last4=Rubenstein |first4=D. I. |last5=Chan-Golston |first5=A. C. |last6=Li |first6=E. |last7=Smith |first7=T. B. |year=2015 |title=How the zebra got its stripes: a problem with too many solutions |journal=Royal Society Open Science |volume=2 |issue=1 |page=140452 |doi=10.1098/rsos.140452|pmid=26064590 |pmc=4448797 |bibcode=2015RSOS....240452L }}</ref><ref name="fitness" /> although one problem with this is that the mountain zebra lives in similar environments and has a bold striping pattern.<ref name="fitness">{{Cite journal |last1=Ruxton |first1=G. D. |title=The possible fitness benefits of striped coat coloration for zebra |doi=10.1046/j.1365-2907.2002.00108.x |journal=Mammal Review |volume=32 |issue=4 |pages=237–244 |year=2002 }}</ref> A 2014 study strongly supported the biting-fly hypothesis, and the quweggaquagga appears to have lived in areas with lesser amounts of fly activity than other zebras.<ref>{{Cite journal |doi=10.1038/ncomms4535 |pmid=24691390 |title=The function of zebra stripes |journal=Nature Communications |volume=5 |pages=3535 |year=2014 |last1=Caro |first1=T. |author1-link=Tim Caro |last2=Izzo |first2=A. |last3=Reiner |first3=R. C. |last4=Walker |first4=H. |last5=Stankowich |first5=T. |bibcode=2014NatCo...5.3535C|doi-access=free }}</ref>

A 2020 study suggested that the [[sexual dimorphism]] in size, with quweggaquagga mares being larger than stallions, could be due to the cold and droughts that affects the Karoo plateau, conditions that were even more severe in prehistoric times, such as during [[ice age]]s (other plains zebras live in warmer areas). Isolation, cold, and aridity could thereby have affected quweggaquagga evolution, including coat colour and size dimorphism. Since plains zebra mares are pregnant or lactate for much of their lives, larger size could have been a selective advantage for quweggaquagga mares, as they would therefore have more food reserves when food was scarce. Dimorphism and coat colour could also have evolved through [[genetic drift]] due to isolation, but these influences are not mutually exclusive, and could have worked together.<ref name="Dimorphism"/>

quweggasQuaggas have been identified in [[cave art]] attributed to the indigenous [[San people]] of Southern Africa.<ref>{{Cite journal |last1=Ouzman |first1=S. |last2=Taçon |first2=P. S. C. |last3=Mulvaney |first3=K. |last4=Fullager |first4=R. |title=Extraordinary Engraved Bird Track from North Australia: Extinct Fauna, Dreaming Being and/or Aesthetic Masterpiece? |doi=10.1017/S0959774302000057 |journal=Cambridge Archaeological Journal |volume=12 |pages=103–112 |year=2002 |s2cid=162218561 }}</ref> As it was easy to find and kill, the quweggaquagga was hunted by early Dutch settlers and later by [[Afrikaners]] to provide meat or for their skins. The skins were traded or exploited. The quweggaquagga was probably vulnerable to extinction due to its restricted range.<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|url-access=limited|publisher=Cambridge University Press|page=[https://archive.org/details/conservingliving00wedd/page/n63 46]|isbn=978-0-521-78812-0}}</ref> Local farmers used them as guards for their livestock, as they were likely to attack intruders.<ref name="Weddell" /> quweggasQuaggas were said to be lively and highly strung, especially the stallions. quweggasQuaggas were brought to European zoos, and an attempt at captive breeding was made at London Zoo, but this was halted when a lone stallion killed itself by bashing itself against a wall after losing its temper.<ref name="Piper2009">{{Cite book |last=Piper |first=R. |title=Extinct animals: an encyclopedia of species that have disappeared during human history |url=https://books.google.com/books?id=k2tFAQAAIAAJ |access-date=23 June 2013 |date=20 March 2009 |publisher=Greenwood Press |isbn=978-0-313-34987-4 |pages=33–36}}</ref> On the other hand, captive quweggasquaggas in European zoos were said to be tamer and more docile than Burchell's zebra.<ref name="Walker" /> One specimen was reported to have lived in captivity for 21 years and 4 months, dying in 1872.<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>

The quweggaquagga was long regarded a suitable candidate for domestication, as it counted as the most docile of the zebras. The Dutch colonists in South Africa had considered this possibility, because their imported work horses did not perform very well in the extreme climate and regularly fell prey to the feared [[African horse sickness]].<ref name="unnatural">{{Cite book |last1=van Grouw |first1=K. |title=Unnatural Selection |date=2018 |publisher=Princeton University Press |isbn=9780691157061 |pages=84–85 }}</ref><ref>{{Cite journal |last1=Ridgeway |first1=W. |title=The Origin and Influence of the Thoroughbred Horse |journal = Cambridge Biological Series | publisher=Cambridge University Press |date=1905 |location=Cambridge |pages=76–78 |doi=10.5962/bhl.title.24156 |url=https://www.biodiversitylibrary.org/item/65396#page/96/mode/1up|doi-access=free }}</ref> In 1843, the English naturalist [[Charles Hamilton Smith]] wrote that the quweggaquagga was 'unquestionably best calculated for domestication, both as regards strength and docility'. Some mentions have been given of tame or domesticated quweggasquaggas in South Africa. In Europe, two stallions were used to drive a [[Phaeton (carriage)|phaeton]] by the sheriff of London in the early 19th&nbsp;century.<ref name="Smith">{{Cite book |last1=Smith |first1=C. H. |title=The Natural History of Horses |publisher=W. H. Lizar |date=1841 |location=Edinburgh |page=331 |doi=10.5962/bhl.title.21334 |url=https://www.biodiversitylibrary.org/item/61546#page/387/mode/1up}}</ref><ref>{{Cite journal |last1=L. |first1=R. |title=Some Animals Exterminated During the Nineteenth Century |journal=Nature |date=1901 |volume=63 |issue=1628 |pages=252–254 |doi=10.1038/063252e0|bibcode=1901Natur..63..252L |doi-access=free }}</ref>

In an attempt at domesticating the quweggaquagga, the British lord [[George Douglas, 16th Earl of Morton]] obtained a single male which he bred with a female horse of partial [[Arabian horse|Arabian]] ancestry. This produced a female [[zebroid|hybrid]] with stripes on its back and legs. [[Lord Morton's mare]] was sold and was subsequently bred with a black stallion, resulting in offspring that again had zebra stripes. An account of this was published in 1820 by the [[Royal Society]].<ref name="Morton">{{Cite journal |last1=Morton |first1=Earl of |title=A Communication of a Singular Fact in Natural History |journal=Philosophical Transactions of the Royal Society of London |date=1821 |volume=111 |pages=20–22 |url=https://archive.org/details/philtrans09227786 |jstor=107600 |issn=0261-0523}}</ref><ref>{{Cite book|author=Birkhead, T. R.|year=2003|title=A Brand New Bird: How Two Amateur Scientists Created the First Genetically Engineered Animal|publisher=Basic Books|page=145|isbn=978-0-465-00665-6}}</ref> It is unknown what happened to the hybrid mare itself. This led to new ideas on [[telegony (pregnancy)|telegony]], referred to as [[pangenesis]] by the British naturalist [[Charles Darwin]].<ref name="Heywood">{{Cite journal | last1 = Heywood | first1 = P. | title = The quagga and science: What does the future hold for this extinct zebra? | doi = 10.1353/pbm.2013.0008 | journal = Perspectives in Biology and Medicine | volume = 56 | issue = 1 | pages = 53–64 | year = 2013 | pmid = 23748526| s2cid = 7991775 }}</ref> At the close of the 19th&nbsp;century, the Scottish zoologist [[James Cossar Ewart]] argued against these ideas and proved, with several cross-breeding experiments, that zebra stripes could appear as an [[atavistic]] trait at any time.<ref name="Penycuik">{{Cite book |last1=Ewart |first1=J. C. |title=The Penycuik Experiments |date=1899 |publisher=A. and C. Black |location=London |pages=55–161 |url=https://www.biodiversitylibrary.org/item/67436#page/153/mode/1up |doi=10.5962/bhl.title.25674}}</ref><ref>{{Cite journalbook |last1=Weeks |first1=D. E. |titlechapter=Newton Morton's influence on genetics: The Morton number |journaltitle=Advances in Genetics <!--|series=Advances in Genetics--> |date=2001 |volume=42 |pages=7–10 |doi=10.1016/s0065-2660(01)42011-6|pmid=11037310 |isbn=9780120176427 }}</ref>

There are 23 known stuffed and mounted quweggaquagga specimens throughout the world, including a juvenile, two foals, and a foetus. In addition, a mounted head and neck, a foot, seven complete skeletons, and samples of various tissues remain. A 24th mounted specimen was destroyed in [[Königsberg]], Germany, during [[World War&nbsp;II]], and various skeletons and bones have also been lost.<ref>{{Cite journal | last = Rau | first = R. E. | title = Revised list of the preserved material of the extinct Cape colony quagga, ''Equus quagga quagga'' (Gmelin) | journal = Annals of the South African Museum. Annale van die Suid-Afrikaanse Museum | volume = 65 | url = https://www.biodiversitylibrary.org/page/40928201#page/67/mode/1up | pages = 41–87 | year = 1974}}</ref><ref>{{Cite journal | last = Rau | first = R. E. | title = Additions to the revised list of preserved material of the extinct Cape Colony quagga and notes on the relationship and distribution of southern plains zebras | journal = Annals of the South African Museum | volume = 77 | year = 1978 | pages = 27–45 | issn = 0303-2515 | url = https://www.biodiversitylibrary.org/page/40680486#page/5/mode/1up }}</ref>

The quweggaquagga had disappeared from much of its range by the 1850s. The last population in the wild, in the [[Orange Free State]], was [[extirpated]] in the late 1870s.<ref name="Walker"/> The last known wild quweggaquagga died in 1878.<ref name="Weddell"/> The specimen in London died in 1872 and the one in Berlin in 1875. The last captive quweggaquagga, a female in Amsterdam's [[Natura Artis Magistra]] zoo, lived there from 9&nbsp;May 1867 until it died on 12&nbsp;August 1883, but its origin and cause of death are unclear.<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 | issue = 8 | year = 1959 | pages = 197–200 | hdl=10520/AJA00382353_1382 |hdl-access=free | url = https://journals.co.za/doi/pdf/10.10520/AJA00382353_1382}}</ref> Its death was not recognised as signifying the extinction of its kind at the time, and the zoo requested another specimen; hunters believed it could still be found "closer to the interior" in the Cape Colony. Since locals used the term quweggaquagga to refer to all zebras, this may have led to the confusion. The extinction of the quweggaquagga was internationally accepted by the 1900 [[Convention for the Preservation of Wild Animals, Birds and Fish in Africa]]. The last specimen was featured on a Dutch stamp in 1988.<ref>{{Cite journal|last1=De Vos|first1=R.|title=Stripes faded, barking silenced: remembering quagga|journal=Animal Studies Journal|date=2014|volume=3|issue=1|url=http://ro.uow.edu.au/asj/vol3/iss1/4/|issn=2201-3008}}</ref> The specimen itself was mounted and is kept in the collection of [[Naturalis Biodiversity Center]] in [[Leiden]]. It has been on display for special occasions.<ref>{{Cite web|title=Naturalis Topstukken|url=http://topstukken.naturalis.nl/object/allerlaatste-quagga|access-date=8 April 2021|website=topstukken.naturalis.nl|language=nl}}</ref>

In 1889, the naturalist [[Henry Anderson Bryden|Henry Bryden]] wrote: "That an animal so beautiful, so capable of domestication and use, and to be found not long since in so great abundance, should have been allowed to be swept from the face of the earth, is surely a disgrace to our latter-day civilization."<ref>{{Cite book|title=Kloof and Karoo|last=Bryden|first=H.|publisher=Longmans, Green and Co|year=1889|asin=B00CNE0EZC|location=London|pages=393–403|url=https://archive.org/details/kloofkarroosport00brydiala/page/392/mode/2up?q=quagga}}</ref>

After the very close relationship between the quweggaquagga and extant plains zebras was discovered, Rau started the quweggaQuagga Project in 1987 in South Africa to create a quweggaquagga-like zebra population by [[selectively breeding]] for a reduced stripe pattern from plains zebra stock, with the eventual aim of introducing them to the quweggaquagga's former range. To differentiate between the quweggaquagga and the zebras of the project, they refer to it as "Rau quweggasquaggas".<ref name="Heywood"/> The founding population consisted of 19 individuals from Namibia and South Africa, chosen because they had reduced striping on the rear body and legs. The first foal of the project was born in 1988. Once a sufficiently quweggaquagga-like population has been created, participants in the project plan to release them in the Western Cape.<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><ref>{{Cite news |url=http://www.cnn.com/2016/01/25/africa/quagga-project-zebra-conservation-extinct-south-africa/ |title=Zebra cousin became extinct 100&nbsp;years ago. Now, it's back |website=CNN|last1=Page|first1=T.|last2=Hancock|first2=C.|date=25 January 2016}}</ref>

Introduction of these quweggaquagga-like zebras could be part of a comprehensive restoration programme, including such ongoing efforts as eradication of non-native trees. quweggasQuaggas, [[wildebeest]], and [[ostrich]]es, which occurred together during historical times in a mutually beneficial association, could be kept together in areas where the indigenous vegetation has to be maintained by grazing. In early 2006, the third- and fourth-generation animals produced by the project were considered looking much like the depictions and preserved specimens of the quweggaquagga. This type of selective breeding is called [[breeding back]]. The practice is controversial, since the resulting zebras will resemble the quweggasquaggas only in external appearance, but will be genetically different. The technology to use recovered DNA for [[cloning]] has not yet been developed.<ref name="Max" /><ref>{{Cite book |last1=Freeman |first1=C. |title=Leonardo¿'s Choice |chapter=Ending Extinction: The Quagga, the Thylacine, and the "Smart Human" |doi=10.1007/978-90-481-2479-4_13 |editor-last1=Gigliotti|editor-first1=C. |pages=235–256 |year=2009 |isbn=978-90-481-2478-7}}</ref>