Editing Quagga (section)

==Taxonomy==
It has been historically suggested that the name ''quagga'' is derived from the [[Khoikhoi]] word for ''[[zebra]]'' ([[cf.]] [[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''.]]

The quagga was originally classified as a distinct [[species]], ''Equus quagga'', 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 quagga 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 quagga in relation to the plains zebra. The  British zoologist [[Reginald Innes Pocock]] in 1902 was perhaps the first to suggest that the quagga was a [[subspecies]] of the plains zebra. As the quagga was scientifically described and named before the plains zebra, the [[trinomial name]] for the quagga becomes ''E. quagga quagga'' under this scheme, and the other subspecies of the plains zebra are placed under ''E. quagga'', as well.<ref name="Azzaroli"/>

Historically, quagga taxonomy was further complicated because the extinct southernmost population of [[Burchell's zebra]] (''Equus quagga 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 quagga 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 quagga, 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 ''Quagga'' 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 quagga'' 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> Quagga 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 quagga 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"/>

===Evolution===

The quagga 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 "quagga" referred to all zebras.<ref name="skinner" /> Fossil skulls of ''[[Equus mauritanicus]]'' from Algeria have been claimed to show affinities with the quagga 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>

{{multiple image
| align = right
| perrow=2/2
| total_width = 350
| header = 
| image1 = Berlin Quagga.jpg
| alt1 =
| image2 = Bamberg Naturkundemuseum Quagga 17RM1912.jpg
| alt2=
| image3 = Equus quagga quagga12.JPG
| image4 = Quagga Naturhistorisches Museum Basel 27102013 3.jpg
|footer = [[Taxidermy]] specimens in Naturkunde-Museum, Bamberg, [[Natural History Museum, Berlin]], [[Museo Civico di Storia Naturale di Milano]], and [[Naturhistorisches Museum, Basel]] (the two above have been sampled for [[DNA]])
}}

The quagga 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 quagga was more closely related to zebras than to horses,<ref name="smithsonian"/> with the quagga 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 quagga 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 quagga 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 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 [[genetic study]] published in 2005 confirmed the subspecific status of the quagga. It showed that the quagga 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 quagga 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>

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

A 2018 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>