Editing Quagga (section)

==Relationship with humans==

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|image1 = Agasse Quagga.jpg
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|image2 = Lord Morton's hybrid mare.jpg
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|footer = Paintings of [[Lord Morton]]'s quagga [[stallion]] (left) and [[Lord Morton's mare]], its [[zebroid|hybrid]] offspring with a horse mare, by [[Jacques-Laurent Agasse]], 1821

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Quaggas 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 quagga was hunted by early Dutch settlers and later by [[Afrikaners]] to provide meat or for their skins. The skins were traded or exploited. The quagga 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" /> Quaggas were said to be lively and highly strung, especially the stallions. Quaggas 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 quaggas 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 quagga 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 quagga was 'unquestionably best calculated for domestication, both as regards strength and docility'. Some mentions have been given of tame or domesticated quaggas 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 quagga, 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 book |last1=Weeks |first1=D. E.   |chapter=Newton Morton's influence on genetics: The Morton number |title=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 quagga 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>

===Extinction===

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|image1 = Quagga exhibit Leiden.jpg
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| caption1 = The last known quagga and a [[great auk]] (another famous case of human-caused extinction), in [[Naturalis]], [[Leiden]]

|image2 = Em - Equus quagga quagga - GMZ 1.jpg
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| caption2 = One of seven known skeletons, at [[Grant Museum]]
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The quagga 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 quagga died in 1878.<ref name="Weddell"/> The specimen in London died in 1872 and the one in Berlin in 1875. 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, 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 quagga to refer to all zebras, this may have led to the confusion. The extinction of the quagga 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>

===Breeding back project===
{{main|Quagga Project}}

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|image1 = Plains Zebra (Equus quagga burchellii) mare, showing the disappearance of stripes characteristic of the "Quagga" proper (now extinct) ... (50220991353).jpg
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|image2 = Buffaloes and Plains Zebras (Equus quagga burchellii) one of them presenting an extreme reduction of the "zebra" pattern ...... (32309547213).jpg
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|footer = [[Quagga Project]] zebras along with regular [[plains zebra]]s (right) in [[Mokala National Park]], South Africa
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After the very close relationship between the quagga and extant plains zebras was discovered, Rau started the Quagga Project in 1987 in South Africa to create a quagga-like zebra population by [[selectively breeding]] for a reduced stripe pattern from plains zebra stock, with the eventual aim of introducing them to the quagga's former range. To differentiate between the quagga and the zebras of the project, they refer to it as "Rau quaggas".<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 quagga-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 quagga-like zebras could be part of a comprehensive restoration programme, including such ongoing efforts as eradication of non-native trees. Quaggas, [[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 quagga. This type of selective breeding is called [[breeding back]]. The practice is controversial, since the resulting zebras will resemble the quaggas 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>