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The quagga ( or ) (''Equus quagga quagga'') is a
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
of the
plains zebra The plains zebra (''Equus quagga'', formerly ''Equus burchellii''), also known as the common zebra, is the most common and geographically widespread species of zebra. Its range is fragmented, but spans much of southern and eastern Africa south o ...
that was
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
to
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
until it was hunted to
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the Endling, last individual of the species, although the Functional ext ...
in the late 19th 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 or
ecotype In evolutionary ecology, an ecotype,Greek: ''οίκος'' = home and ''τύπος'' = type, coined by Göte Turesson in 1922 sometimes called ecospecies, describes a genetically distinct geographic variety, population, or race within a species ...
of the species. The quagga is believed to have been around long and tall at the shoulder. It was distinguished from other
zebra Zebras (, ) (subgenus ''Hippotigris'') are African equines with distinctive black-and-white striped coats. There are three living species: the Grévy's zebra (''Equus grevyi''), plains zebra (''E. quagga''), and the mountain zebra (''E. zebr ...
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 Burchell's zebra (''Equus quagga burchellii'') is a southern subspecies of the plains zebra. It is named after the British explorer and naturalist William John Burchell. Common names include bontequagga, Damaraland zebra, and Zululand zebra (Gray, ...
. They were once found in great numbers in the
Karoo The Karoo ( ; from the Afrikaans borrowing of the South Khoekhoe !Orakobab or Khoemana word ''ǃ’Aukarob'' "Hardveld") is a semi-desert natural region of South Africa. No exact definition of what constitutes the Karoo is available, so its ex ...
of
Cape Province The Province of the Cape of Good Hope ( af, Provinsie Kaap die Goeie Hoop), commonly referred to as the Cape Province ( af, Kaapprovinsie) and colloquially as The Cape ( af, Die Kaap), was a province in the Union of South Africa and subsequen ...
and the southern part of the
Orange Free State The Orange Free State ( nl, Oranje Vrijstaat; af, Oranje-Vrystaat;) was an independent Boer sovereign republic under British suzerainty in Southern Africa during the second half of the 19th century, which ceased to exist after it was defeat ...
in South Africa. After the European settlement of South Africa began, the quagga 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 quagga was
extinct in the wild A species that is extinct in the wild (EW) is one that has been categorized by the International Union for Conservation of Nature as known only by living members kept in captivity or as a naturalized population outside its historic range due t ...
by 1878. The last captive specimen died in Amsterdam on 12 August 1883. Only one quagga was ever photographed alive, and only 23 skins exist today. In 1984, the quagga was the first extinct animal whose DNA was analysed. The
Quagga Project The Quagga Project is an attempt by a group in South Africa to use selective breeding to achieve a breeding lineage of Burchell's zebra (''Equus quagga burchellii'') which visually resemble the extinct quagga (''Equus quagga quagga''). History ...
is trying to recreate the
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (biology), morphology or physical form and structure, its Developmental biology, developmental proc ...
of hair coat pattern by selectively breeding the genetically closest subspecies, which is Burchell's zebra.


Taxonomy

It has been historically suggested that the name ''quagga'' is derived from the
Khoikhoi Khoekhoen (singular Khoekhoe) (or Khoikhoi in the former orthography; formerly also '' Hottentots''"Hottentot, n. and adj." ''OED Online'', Oxford University Press, March 2018, www.oed.com/view/Entry/88829. Accessed 13 May 2018. Citing G. S. ...
word for ''
zebra Zebras (, ) (subgenus ''Hippotigris'') are African equines with distinctive black-and-white striped coats. There are three living species: the Grévy's zebra (''Equus grevyi''), plains zebra (''E. quagga''), and the mountain zebra (''E. zebr ...
'' (
cf. The abbreviation ''cf.'' (short for the la, confer/conferatur, both meaning "compare") is used in writing to refer the reader to other material to make a comparison with the topic being discussed. Style guides recommend that ''cf.'' be used onl ...
Tshwa 'zebra'), thereby being an
onomatopoeic Onomatopoeia is the process of creating a word that phonetically imitates, resembles, or suggests the sound that it describes. Such a word itself is also called an onomatopoeia. Common onomatopoeias include animal noises such as ''oink'', ''m ...
word, resembling the quagga's call, variously transcribed as "kwa-ha-ha", "kwahaah", or "oug-ga". The name is still used colloquially for the
plains zebra The plains zebra (''Equus quagga'', formerly ''Equus burchellii''), also known as the common zebra, is the most common and geographically widespread species of zebra. Its range is fragmented, but spans much of southern and eastern Africa south o ...
. The quagga was originally classified as a distinct
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
, ''Equus quagga'', in 1778 by Dutch naturalist
Pieter Boddaert Pieter Boddaert (1730 – 6 May 1795) was a Dutch physician and naturalist. Early life, family and education Boddaert was the son of a Middelburg jurist and poet by the same name (1694–1760). The younger Pieter obtained his M.D. at the Unive ...
. Traditionally, the quagga and the other plains and mountain zebras were placed in the
subgenus In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between ...
''
Hippotigris Zebras (, ) (subgenus ''Hippotigris'') are African equines with distinctive black-and-white striped coats. There are three living species: the Grévy's zebra (''Equus grevyi''), plains zebra (''E. quagga''), and the mountain zebra (''E. zebr ...
''. Much debate has occurred over the status of the quagga in relation to the plains zebra. The British zoologist
Reginald Innes Pocock Reginald Innes Pocock Fellow of the Royal Society, F.R.S. (4 March 1863 – 9 August 1947) was a British zoologist. Pocock was born in Clifton, Bristol, the fourth son of Rev. Nicholas Pocock (historian), Nicholas Pocock and Edith Prichard. He ...
in 1902 was perhaps the first to suggest that the quagga was a
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
of the plains zebra. As the quagga was scientifically described and named before the plains zebra, the
trinomial name In biology, trinomial nomenclature refers to names for taxa below the rank of species. These names have three parts. The usage is different in zoology and botany. In zoology In zoological nomenclature, a trinomen (), trinominal name, or ternar ...
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. Historically, quagga taxonomy was further complicated because the extinct southernmost population of
Burchell's zebra Burchell's zebra (''Equus quagga burchellii'') is a southern subspecies of the plains zebra. It is named after the British explorer and naturalist William John Burchell. Common names include bontequagga, Damaraland zebra, and Zululand zebra (Gray, ...
(''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 In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
. The extinct population was long thought very close to the quagga, since it also showed limited striping on its hind parts. As an example of this, Shortridge placed the two in the now disused subgenus ''Quagga'' in 1934. Most experts now suggest that the two subspecies represent two ends of a cline. Different subspecies of plains zebras were recognised as members of ''Equus quagga'' by early researchers, though much confusion existed over which species were valid. 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. Some subspecies and even species, such as ''E. q. danielli'' and ''Hippotigris isabellinus'', were based only on illustrations (iconotypes) of aberrant quagga specimens. One craniometric study from 1980 seemed to confirm its affiliation with the
horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million yea ...
(''Equus ferus caballus''), but early morphological studies have been noted as being erroneous. Studying skeletons from stuffed specimens can be problematical, as early taxidermists sometimes used
donkey The domestic donkey is a hoofed mammal in the family Equidae, the same family as the horse. It derives from the African wild ass, ''Equus africanus'', and may be classified either as a subspecies thereof, ''Equus africanus asinus'', or as ...
and horse skulls inside their mounts when the originals were unavailable.


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. 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. The quagga was the first extinct animal to have its DNA analysed, and this 1984 study launched the field of
ancient DNA Ancient DNA (aDNA) is DNA isolated from ancient specimens. Due to degradation processes (including cross-linking, deamination and fragmentation) ancient DNA is more degraded in comparison with contemporary genetic material. Even under the bes ...
analysis. It confirmed that the quagga was more closely related to zebras than to horses, with the quagga and
mountain zebra The mountain zebra (''Equus zebra'') is a zebra species in the family Equidae, native to southwestern Africa. There are two subspecies, the Cape mountain zebra (''E. z. zebra'') found in South Africa and Hartmann's mountain zebra (''E. z. hartman ...
(''Equus zebra'') sharing an ancestor 3–4 million years ago. An
immunological Immunology is a branch of medicineImmunology for Medical Students, Roderick Nairn, Matthew Helbert, Mosby, 2007 and biology that covers the medical study of immune systems in humans, animals, plants and sapient species. In such we can see the ...
study published the following year found the quagga to be closest to the plains zebra. A 1987 study suggested that the
mtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
of the quagga diverged at a range of roughly 2 percent per million years, similar to other mammal species, and again confirmed the close relation to the plains zebra. 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. A 2004 study of skins and skulls instead suggested that the quagga was not a distinct species, but a subspecies of the plains zebra. In spite of these findings, many authors subsequently kept the plains zebra and the quagga as separate species. A
genetic study Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar working i ...
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 The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
, 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 Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, ...
s diverged into separate species and subspecies during this period, as well, probably because of the same climate shift. The simplified
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
below is based on the 2005 analysis (some taxa shared
haplotype A haplotype ( haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material ( DNA) which is inherited from two parents. Normally these organisms have their DNA o ...
s and could, therefore, not be differentiated): 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 }), is a landlocked country in East Africa. The country is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the south by Tanzania. The ...
. Instead, the study supported a north–south genetic continuum for plains zebras, with the Ugandan population being the most distinct. Zebras from
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
appear to be the closest genetically to the quagga.


Description

The quagga is believed to have been long and tall at the shoulder. Based on measurements of skins, mares were significantly longer and slightly taller than stallions, whereas the stallions of extant zebras are the largest. Its coat pattern was unique among
equid Equidae (sometimes known as the horse family) is the taxonomic family of horses and related animals, including the extant horses, asses, and zebras, and many other species known only from fossils. All extant species are in the genus '' Equus' ...
s: zebra-like in the front but more like a horse in the rear. 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, 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. It also had a broad dark dorsal stripe on its back. It had a standing mane with brown and white stripes. The only quagga to have been photographed alive was a
mare A mare is an adult female horse or other equine. In most cases, a mare is a female horse over the age of three, and a filly is a female horse three and younger. In Thoroughbred horse racing, a mare is defined as a female horse more than fo ...
at the
Zoological Society of London The Zoological Society of London (ZSL) is a charity devoted to the worldwide conservation of animals and their habitats. It was founded in 1826. Since 1828, it has maintained the London Zoo, and since 1931 Whipsnade Park. History On 29 ...
's
Zoo A zoo (short for zoological garden; also called an animal park or menagerie) is a facility in which animals are kept within enclosures for public exhibition and often bred for conservation purposes. The term ''zoological garden'' refers to z ...
. Five photographs of this specimen are known, taken between 1863 and 1870. On the basis of photographs and written descriptions, many observers suggest that the stripes on the quagga were light on a dark background, unlike other zebras. The German naturalist Reinhold Rau, pioneer of the
Quagga Project The Quagga Project is an attempt by a group in South Africa to use selective breeding to achieve a breeding lineage of Burchell's zebra (''Equus quagga burchellii'') which visually resemble the extinct quagga (''Equus quagga quagga''). History ...
, claimed that this is an
optical illusion Within visual perception, an optical illusion (also called a visual illusion) is an illusion caused by the visual system and characterized by a visual perception, percept that arguably appears to differ from reality. Illusions come in a wide v ...
: that the base colour is a creamy white and that the stripes are thick and dark. Living in the very southern end of the plains zebra's range, the quagga had a thick winter coat that moulted each year. Its skull was described as having a straight profile and a
concave Concave or concavity may refer to: Science and technology * Concave lens * Concave mirror Mathematics * Concave function, the negative of a convex function * Concave polygon, a polygon which is not convex * Concave set In geometry, a subset o ...
diastema A diastema (plural diastemata, from Greek διάστημα, space) is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition ...
, and as being relatively broad with a narrow occiput. Like other plains zebras, the quagga did not have a
dewlap A dewlap is a longitudinal flap of skin or similar flesh that hangs beneath the lower jaw or neck of many vertebrates. More loosely, it can be various similar structures in the neck area, such as those caused by a double chin or the submandibul ...
on its neck as the mountain zebra does. The 2004 morphological study found that the skeletal features of the southern Burchell's zebra population and the quagga 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 quaggas and southern Burchell's zebra are so similar that they are impossible to definitely identify as either, since no location data was recorded.


Behaviour and ecology

The quagga was the southernmost distributed plains zebra, mainly living south of the
Orange River The Orange River (from Afrikaans/Dutch: ''Oranjerivier'') is a river in Southern Africa. It is the longest river in South Africa. With a total length of , the Orange River Basin extends from Lesotho into South Africa and Namibia to the north ...
. It was a grazer, and its habitat range was restricted to the grasslands and arid interior scrubland of the
Karoo The Karoo ( ; from the Afrikaans borrowing of the South Khoekhoe !Orakobab or Khoemana word ''ǃ’Aukarob'' "Hardveld") is a semi-desert natural region of South Africa. No exact definition of what constitutes the Karoo is available, so its ex ...
region of
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
, today forming parts of the provinces of
Northern Cape The Northern Cape is the largest and most sparsely populated province of South Africa. It was created in 1994 when the Cape Province was split up. Its capital is Kimberley. It includes the Kalahari Gemsbok National Park, part of the Kgalagadi ...
,
Eastern Cape The Eastern Cape is one of the provinces of South Africa. Its capital is Bhisho, but its two largest cities are East London and Gqeberha. The second largest province in the country (at 168,966 km2) after Northern Cape, it was formed in ...
,
Western Cape The Western Cape is a province of South Africa, situated on the south-western coast of the country. It is the fourth largest of the nine provinces with an area of , and the third most populous, with an estimated 7 million inhabitants in 202 ...
, and the Free State. These areas were known for distinctive flora and fauna and high amounts of
endemism Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found els ...
. Quaggas have been reported gathering into herds of 30–50, and sometimes travelled in a linear fashion. They may have been
sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species s ...
with Burchell's zebra between the
Vaal The Vaal River ( ; Khoemana: ) is the largest tributary of the Orange River in South Africa. The river has its source near Breyten in Mpumalanga province, east of Johannesburg and about north of Ermelo and only about from the Indian Ocean. I ...
and Orange rivers. This is disputed, and there is no evidence that they interbred. It could also have shared a small portion of its range with
Hartmann's mountain zebra Hartmann's mountain zebra (''Equus zebra hartmannae'') is a subspecies of the mountain zebra found in far south-western Angola and western Namibia, easily distinguished from other similar zebra species by its dewlap as well as the lack of strip ...
(''Equus zebra hartmannae''). Little is known about the behaviour of quaggas in the wild, and it is sometimes unclear what exact species of zebra is referred to in old reports. The only source that unequivocally describes the quagga in the Free State is that of the British military engineer and hunter
William Cornwallis Harris Major Sir William Cornwallis Harris (baptised 2 April 1807 – died 9 October 1848) was an English military engineer, artist and hunter. Life and career Early life The son of James Harris of Wittersham, Kent, he entered Addiscombe Militar ...
. His 1840 account reads as follows:
The geographical range of the quagga 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 quaggas 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 A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
and flowers to form a green carpet, spangled with hues the most brilliant and diversified.Sir Cornwallis Harris, quoted in
The practical function of striping in zebras has been debated and it is unclear why the quagga lacked stripes on its hind parts. A
cryptic Cryptic may refer to: In science: * Cryptic species complex, a group of species that are very difficult to distinguish from one another * Crypsis, the ability of animals to blend in to avoid observation * Cryptic era, earliest period of the Earth ...
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 Intra-species recognition is the recognition by a member of a species of a conspecific (another member of the same species). In many species, such recognition is necessary for procreation. Different species may employ different methods, but all ...
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 Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperatur ...
to cool themselves down, and that the quagga lost them due to living in a cooler climate, although one problem with this is that the mountain zebra lives in similar environments and has a bold striping pattern. A 2014 study strongly supported the biting-fly hypothesis, and the quagga appears to have lived in areas with lesser amounts of fly activity than other zebras. A 2020 study suggested that the
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
in size, with quagga 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 An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gre ...
s (other plains zebras live in warmer areas). Isolation, cold, and aridity could thereby have affected quagga 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 quagga mares, as they would therefore have more food reserves when food was scarce. Dimorphism and coat colour could also have evolved through
genetic drift Genetic drift, also known as allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene variants to disappear completely and there ...
due to isolation, but these influences are not mutually exclusive, and could have worked together.


Relationship with humans

Quaggas have been identified in
cave art In archaeology, Cave paintings are a type of parietal art (which category also includes petroglyphs, or engravings), found on the wall or ceilings of caves. The term usually implies prehistoric origin, and the oldest known are more than 40,000 ye ...
attributed to the indigenous
San people The San peoples (also Saan), or Bushmen, are members of various Khoe, Tuu, or Kxʼa-speaking indigenous hunter-gatherer cultures that are the first cultures of Southern Africa, and whose territories span Botswana, Namibia, Angola, Zam ...
of Southern Africa. As it was easy to find and kill, the quagga was hunted by early Dutch settlers and later by
Afrikaners Afrikaners () are a South African ethnic group descended from predominantly Dutch settlers first arriving at the Cape of Good Hope in the 17th and 18th centuries.Entry: Cape Colony. ''Encyclopædia Britannica Volume 4 Part 2: Brain to Cas ...
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. Local farmers used them as guards for their livestock, as they were likely to attack intruders. Quaggas were said to be lively and highly strung, especially the stallions. Quaggas were brought to European zoos, and an attempt at captive breeding at London Zoo, but was halted when a lone stallion killed itself by bashing itself against a wall after losing its temper. On the other hand, captive quaggas in European zoos were said to be tamer and more docile than Burchell's zebra. One specimen was reported to have lived in captivity for 21 years and 4 months, dying in 1872. 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 African horse sickness (AHS) is a highly infectious and deadly disease caused by ''African horse sickness virus''. It commonly affects horses, mules, and donkeys. It is caused by a virus of the genus ''Orbivirus'' belonging to the family ' ...
. In 1843, the English naturalist
Charles Hamilton Smith Lieutenant-Colonel Charles Hamilton Smith, KH, KW, FRS, FLS, (26 December 1776 in East Flanders, in the United Provinces of the Netherlands – 21 September 1859 in Plymouth) was an English artist, naturalist, antiquary, illustrator, soldie ...
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 by the sheriff of London in the early 19th century. In an attempt at domesticating the quagga, the British lord
George Douglas, 16th Earl of Morton George Douglas, 16th Earl of Morton, KT, FRS, FRSE, FSA (3 April 1761 – 17 July 1827) was a British nobleman. Life He was the son of Sholto Douglas, 15th Earl of Morton, and Katherine Hamilton. He succeeded to the title Earl of Morton in 17 ...
obtained a single male which he bred with a female horse of partial
Arabian The Arabian Peninsula, (; ar, شِبْهُ الْجَزِيرَةِ الْعَرَبِيَّة, , "Arabian Peninsula" or , , "Island of the Arabs") or Arabia, is a peninsula of Western Asia, situated northeast of Africa on the Arabian Plat ...
ancestry. This produced a female hybrid with stripes on its back and legs.
Lord Morton's mare Lord Morton’s mare was an equid hybrid and once an often-noticed example in the history of evolutionary theory. In 1820, George Douglas, 16th Earl of Morton, F.R.S., reported to the President of the Royal Society that, being desirous of dome ...
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 The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society and the United Kingdom's national academy of sciences. The society fulfils a number of roles: promoting science and its benefits, re ...
. It is unknown what happened to the hybrid mare itself. This led to new ideas on
telegony The ''Telegony'' (Greek: , ''Tēlegoneia''; la, Telegonia) is a lost ancient Greek epic poem about Telegonus, son of Odysseus by Circe. His name ("born far away") is indicative of his birth on Aeaea, far from Odysseus' home of Ithaca. It was p ...
, referred to as
pangenesis Pangenesis was Charles Darwin's hypothetical mechanism for heredity, in which he proposed that each part of the body continually emitted its own type of small organic particles called gemmules that aggregated in the gonads, contributing herita ...
by the British naturalist
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
. At the close of the 19th century, the Scottish zoologist
James Cossar Ewart James Cossar Ewart FRS FRSE (26 November 1851 – 31 December 1933) was a Scottish zoologist. He performed breeding experiments with horses and zebras which disproved earlier theories of heredity. Life Ewart was born in Penicuik, Midloth ...
argued against these ideas and proved, with several cross-breeding experiments, that zebra stripes could appear as an
atavistic In biology, an atavism is a modification of a biological structure whereby an ancestral genetic trait reappears after having been lost through evolutionary change in previous generations. Atavisms can occur in several ways; one of which is when ...
trait at any time. 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 Königsberg (, ) was the historic Prussian city that is now Kaliningrad, Russia. Königsberg was founded in 1255 on the site of the ancient Old Prussian settlement ''Twangste'' by the Teutonic Knights during the Northern Crusades, and was ...
, Germany, during
World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposi ...
, and various skeletons and bones have also been lost.


Extinction

The quagga had disappeared from much of its range by the 1850s. The last population in the wild, in the
Orange Free State The Orange Free State ( nl, Oranje Vrijstaat; af, Oranje-Vrystaat;) was an independent Boer sovereign republic under British suzerainty in Southern Africa during the second half of the 19th century, which ceased to exist after it was defeat ...
, was
extirpated Local extinction, also known as extirpation, refers to a species (or other taxon) of plant or animal that ceases to exist in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinct ...
in the late 1870s. The last known wild quagga died in 1878. 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 May 1867 until it died on 12 August 1883, but its origin and cause of death are unclear. 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 Convention for the Preservation of Wild Animals, Birds and Fish in Africa (also known as the London Convention of 1900) is a multilateral treaty on wildlife preservation that was signed by the European colonial powers in London in 1900. Alth ...
. The last specimen was featured on a Dutch stamp in 1988. The specimen itself was mounted and is kept in the collection of
Naturalis Biodiversity Center Naturalis Biodiversity Center ( nl, Nederlands Centrum voor Biodiversiteit Naturalis) is a national museum of natural history and a research center on biodiversity in Leiden, Netherlands. It was named the European Museum of the Year 2021. ...
in
Leiden Leiden (; in English and archaic Dutch also Leyden) is a city and municipality in the province of South Holland, Netherlands. The municipality of Leiden has a population of 119,713, but the city forms one densely connected agglomeration w ...
. It has been on display for special occasions. In 1889, the naturalist 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."


Breeding back project

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". 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. 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 Wildebeest ( , , ), also called gnu ( or ), are antelopes of the genus ''Connochaetes'' and native to Eastern and Southern Africa. They belong to the family Bovidae, which includes true antelopes, cattle, goats, sheep, and other even-toe ...
, and
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There ...
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 Breeding back is a form of artificial selection by the deliberate selective breeding of domestic (but not exclusively) animals, in an attempt to achieve an animal breed with a phenotype that resembles a wild type ancestor, usually one that ha ...
. 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 Cloning is the process of producing individual organisms with identical or virtually identical DNA, either by natural or artificial means. In nature, some organisms produce clones through asexual reproduction. In the field of biotechnology, c ...
has not yet been developed.


See also

*
Holocene extinction The Holocene extinction, or Anthropocene extinction, is the ongoing extinction event during the Holocene epoch. The extinctions span numerous families of bacteria, fungi, plants, and animals, including mammals, birds, reptiles, amphibians, fish, ...
* Lists of extinct animals


References


External links

* {{Authority control Zebras Extinct mammals of Africa Mammal extinctions since 1500 Mammals of South Africa Species endangered by use as food Species made extinct by human activities Mammals described in 1785 Subspecies Taxa named by Pieter Boddaert