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Quagga
Live quagga mare in London Zoo, 1870
Conservation status
Extinct (1883) (IUCN 3.1)
[1]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Perissodactyla
Family: Equidae
Genus: Equus
Subgenus: Hippotigris
Species: E. quagga
Subspecies: †E. q. quagga
Trinomial name
†Equus quagga quagga
(Boddaert, 1785)
Quagga
From Wikipedia, the free encyclopedia
The quagga (/ˈkwɑːxɑː/)
[2][3]
(Equus quagga quagga) is an extinct
subspecies of the plains zebra that lived in South Africa until the
nineteenth century. It was long thought to be a distinct species, but
recent genetic studies have shown it to be the southernmost
subspecies of the plains zebra. It is considered particularly close to
Burchell's zebra. Its name is derived from its call, which sounds like
"kwa-ha-ha".
The quagga is believed to have been around 257 cm (8 ft 5 in) long
and 125–135 cm (4 ft 1 in–4 ft 5 in) tall at the shoulder. It was
distinguished from other zebras 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 therefore more horse-like.
The distribution of stripes varied considerably between individuals.
Little is known about its behaviour, but it may have gathered into
herds of 30–50 individuals. Quaggas were said to be wild and lively,
yet were also considered more docile than 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.
Since Dutch settlement of South Africa began, the quagga was
heavily hunted, and it competed with domesticated animals for
forage. While some individuals were taken to zoos in Europe,
breeding programs were not successful. The last wild population
lived in the Orange Free State, and the quagga was extinct in the
wild by 1878. The last captive specimen died in Amsterdam on 12
August 1883. Only one quagga was ever photographed alive and
only 23 skins are preserved today. In 1984, the quagga was the first
extinct animal to have its DNA analysed, and the Quagga Project is
trying to recreate its pelage characteristics by selectively breeding
Burchell's zebras.
Contents
1 Taxonomy
1.1 Evolution
2 Description
3 Behaviour and ecology
4 Decline and extinction
4.1 Breeding back project
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Former range in red
Synonyms 1804 illustration by Samuel Daniell,
which was the basis of the
subspecies E. q. danielli
5 See also
6 References
Taxonomy
The name "quagga" is
derived from the Khoikhoi
word for zebra and is
onomatopoeic, being said
to resemble the quagga's
call, variously transcribed
as "kwa-ha-ha"
[4]
,
"kwahaah"
[2]
, or "oug-
ga".
[5]
The name is still
used colloquially for the
plains zebra.
[4]
The
quagga was originally
classified as a distinct species, Equus quagga, in 1778 by Dutch
naturalist Pieter Boddaert.
[6]
Traditionally, the quagga and the other plains and mountain zebras were placed in the
subgenus Hippotigris.
[7]
There has been much debate over the status of the quagga in relation to the plains zebra. It 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.
[4]
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.
[8]
Quaggas have also been identified in cave art attributed to the San.
[9]
Reginald Innes
Pocock was perhaps the first to suggest that the quagga was a subspecies of plains zebra in 1902. 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 plains zebra are placed under E. quagga as
well.
[8]
Historically, quagga taxonomy was further complicated by the fact that 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.
[7]
As an example of this, Shortridge placed the two in
the now disused subgenus Quagga in 1934.
[10]
Most experts now suggest that the two subspecies represent two
ends of a cline.
[11]
Different subspecies of plains zebra were recognised as members of Equus quagga by early researchers, though
there was much confusion over which species were valid.
[12]
Quagga subspecies were described on the basis of
differences in striping patterns, but these differences were since attributed to individual variation within the same
List
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The mare in London Zoo, 1870
Specimen in the Museum für
Naturkunde, Berlin, which has been
sampled for DNA
Specimen in Naturhistorisches
Museum, Basel
populations.
[13]
Some subspecies and even species, such as E. q. danielli and Hippotigris isabellinus, were only
based on illustrations (iconotypes) of aberrant quagga specimens.
[14][15][16]
Some authors have described the
quagga as a kind of wild horse rather than a zebra, and one craniometric
study from 1980 seemed to confirm its affiliation with the horse (Equus
caballus).
[11]
It has been pointed out that early morphological studies
were erroneous, since using skeletons from stuffed specimens can be
problematical, as early taxidermists sometimes used donkey and horse
skulls inside their mounts when the originals were unavailable.
[17]
Evolution
The quagga was the first
extinct animal to have its DNA
analysed,
[18]
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,
[19]
with the
quagga and mountain zebra (Equus zebra) sharing an ancestor 3–4
million years ago.
[18]
An immunological study published the following
year found the quagga to be closest to the plains zebra.
[20]
A 1987 study
suggested that the mtDNA of the quagga diverged at a range of roughly
2% per million years, similar to other mammal species, and again
confirmed the close relation to the plains zebra.
[21]
Later morphological studies came to conflicting 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.
[19]
A 2004 study of skins and skulls instead suggested
that the quagga was not a distinct species, but a subspecies of the plains zebra.
[7]
In spite of these findings, many
authors subsequently kept the plains zebra and the quagga as separate species.
[4]
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 ungulates diverged into separate
species and subspecies during this period as well, probably because of
the same climate shift. The simplified cladogram below is based on the
2005 analysis (some taxa shared haplotypes and could therefore not be
differentiated):
[19]
Mountain zebra (E. zebra)
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Painting of a stallion in Louis XVI's
menagerie at Versailles by Nicolas
Marechal, 1793
Grévy's zebra (E. grevyi)
Quagga (E. q. quagga)
Damara zebra (E. q. antiquorum)-
Chapman's zebra (E. q. chapmani)
Damara zebra-Chapman's zebra
Grant's zebra (E. q. boehmi)
Grant's zebra
Description
The quagga is believed to have been 257 cm (8 ft 5 in) long and 125–
135 cm (4 ft 1 in–4 ft 5 in) tall at the shoulder.
[11]
Its coat pattern was
unique among equids: zebra-like in the front but more like a horse in the
rear.
[19]
It had brown stripes on the head and neck, brown upper parts
and a white belly, tail and legs. The stripes were darkest on the head and
neck and became gradually lighter 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
individuals 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.
[11]
It
also had a broad dark dorsal stripe on its back. It had a standing mane
with brown and white stripes.
[5]
The only quagga to have been photographed alive was a mare at the Zoological Society of London's Zoo. Five
photographs of this specimen are known, taken between 1863 and 1870.
[22]
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. Reinhold Rau, pioneer of the Quagga Project, claimed that this is an optical illusion: that the base
colour is a creamy white and that the stripes are thick and dark.
[11]
Embryological evidence supports zebras being
dark coloured with white as an addition.
[23]
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The London mare next to a keeper,
1864
Live stallion at the Royal College of
Surgeons, painted by Jacques-Laurent
Agasse in the early 1800s
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 diastema, and as being relatively
broad with a narrow occiput.
[7][24]
Like other plains zebras, the quagga
did not have a dewlap on its neck as the mountain zebra does.
[8]
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 individuals of the extant
Burchell's zebra population still exhibit 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. The female specimens used in the
study were larger than the males on average.
[7]
Behaviour and ecology
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.
[11][25]
These
areas were known for distinctive flora and fauna and high amounts of
endemism.
[24][26]
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.
[11]
The only source that unequivocally describes the quagga in
the Free State is that of the English military engineer and hunter Major Sir
William Cornwallis Harris.
[7]
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 and flowers to form a green carpet, spangled with hues the most brilliant and
diversified.
[27]
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Fifth known photo of the London Zoo
mare, taken in 1863, rediscovered in
1991
Skeleton at the Grant
Museum
Quaggas have been reported gathering into herds of 30–50 individuals
and sometimes travelled in a linear fashion.
[11]
They may have been
sympatric with Burchell's zebra between the Vaal and Orange
rivers.
[7][26]
This is disputed,
[7]
and there is no evidence that they
interbred.
[26]
It could also have shared a small portion of its range with
Hartmann's mountain zebra (Equus zebra hartmannae).
[19]
Quaggas were said to be lively and highly strung, especially the stallions.
During the 1830s, quaggas were used as harness animals for carriages in
London, the males probably being gelded to mitigate their volatile
nature.
[28]
Local farmers used them as guards for their livestock, as they
were likely to attack intruders.
[29]
On the other hand, captive quaggas in
European zoos were said to be tamer and more docile than Burchell's
zebra.
[11]
One specimen was reported to have lived in captivity for 21 years and 4 months, dying in 1872.
[11]
Since the practical function of striping has not been determined for zebras in general, it is unclear why the quagga
lacked stripes on its hind parts. A cryptic function for protection from predators (stripes obscure the individual
zebra in a herd) and tsetse flies (these flies 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 hypothesised that the zebras developed striping patterns as thermoregulation 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.
[30]
A 2014 study supported the biting-fly
hypothesis unambiguously, though it left various aspects of the explanation open to further investigation.
[31]
Decline and extinction
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
used locally. The quagga was probably vulnerable to extinction due to its limited
distribution, and it may have competed with domestic livestock for forage.
[29]
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.
[11]
The
last known wild individual died in 1878.
[29]
Individual quaggas were also captured and shipped to Europe, where they were
displayed in zoos.
[11]
Lord Morton tried to save the animal from extinction by
starting a captive breeding program. He was only able to obtain a single male
which, in desperation, he bred with a female horse. This produced a female
hybrid which bore the zebra 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.
[32]
This lead to new ideas on telegony, referred to as pan-genesis by Charles Darwin.
[25]
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The last known quagga and a great
auk, in Naturalis, Leiden
The last captive specimen, a female in Amsterdam's Natura Artis
Magistra zoo, had lived there from 9 May 1867 until it died on 12
August 1883, but its origin and cause of death were not recorded. The
specimen in London died in 1872 and the one in Berlin in 1875.
[13]
There are 23 known stuffed and mounted quagga specimens throughout
the world. In addition, there is a mounted head and neck, a foot, seven
complete skeletons, and samples of various tissues. A twenty-fourth
mounted specimen was destroyed in Königsberg, Germany, during
World War II.
[33]
Breeding back project
After the very close relationship between the quagga and surviving zebras
was discovered, the Quagga Project was started in 1987 by Reinhold
Rau in South Africa to recreate the quagga by selective breeding from plains zebra stock, with the eventual aim of
reintroducing them to the wild. To differentiate between the previously existing quagga zebras and the ones bred
back into the environment, it has been suggested the new population should be referred to as "Rau quaggas".
[25]
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, it will be released in the Western Cape.
[17]
Introduction of Rau's quaggas could be part of a comprehensive restoration program including such ongoing efforts
as eradication of nonnative trees. Quaggas, wildebeest, and ostriches, 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
reported to look very 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
does not exist.
[2][34]
See also
Lists of extinct animals
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8/12/2014 Quagga - Wikipedia, the free encyclopedia
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Categories: IUCN Red List extinct species Extinct mammals of Africa Mammal extinctions since 1500
Mammals of South Africa Megafauna of Africa Species made extinct by human activities Zebras
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