Quagga: Difference between revisions

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

The '''quagga''' ({{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 quagga quagga''''') is aan extinct [[subspecies]] of the [[plains zebra]] that was [[endemic]] to [[South Africa]] until it was hunted to [[extinction]] in the late 18th19th&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 quagga 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 quagga's behaviour, but it may have gathered into herds of 30–50. Quaggas 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.

It has been historically suggested that the name ''quagga'' 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 quagga'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''.]]

Later morphological studies came to different conclusions. A 1999 analysis of cranial measurements found that the quagga 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 quagga 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 quagga as separate species.<ref name=skinner/>

A 2018 genetic study of plains zebras populations confirmed the quagga as a member of that species. They found no evidence for subspecific differentiation based on morphological differences between southern populations of zebras, including the quagga. Modern plains zebra populations may have originated from southern Africa, and the quagga 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 quagga.<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 quagga 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 quagga 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> Quaggas 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]]

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

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>

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 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>

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>

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>