Quagga

Quagga
Quagga photo.jpg
Quagga mare at London Zoo, 1870, the only specimen photographed alive

Extinct  (1883) ( IUCN 3.1) [1]
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Class: Mammalia
Order: Perissodactyla
Family: Equidae
Genus: Equus
Species: E. quagga
Subspecies: E. q. quagga
Trinomial name
Equus quagga quagga
( Boddaert, 1785)
Quagga range.png
Former range in red
Synonyms

The quagga ( ɑː/ or ə/) [2] [3] [4] (Equus quagga quagga) is an extinct subspecies of plains zebra that lived in South Africa until the 19th century. It was long thought to be a distinct species, but genetic studies have shown it to be the southernmost subspecies of plains zebra. It is considered particularly close to Burchell's zebra. Its name was derived from its call, which sounded 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 the quagga's 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.

After the Dutch settlement of South Africa began, the quagga was heavily hunted as it competed with domesticated animals for forage. While some individuals were taken to zoos in Europe, breeding programs were unsuccessful. 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 the phenotype of hair coat pattern and related characteristics by selectively breeding Burchell's zebras.

Taxonomy

1804 illustration by Samuel Daniell, which was the basis of the supposed subspecies E. q. danielli

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", [5] "kwahaah", [2] or "oug-ga". [6] The name is still used colloquially for the plains zebra. [5] The quagga was originally classified as a distinct species, Equus quagga, in 1778 by Dutch naturalist Pieter Boddaert. [7] Traditionally, the quagga and the other plains and mountain zebras were placed in the subgenus Hippotigris. [8]

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. [5] 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. [9] Quaggas have also been identified in cave art attributed to the San. [10] 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. [9]

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. [8] As an example of this, Shortridge placed the two in the now disused subgenus Quagga in 1934. [11] Most experts now suggest that the two subspecies represent two ends of a cline. [12]

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. [13] 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. [14] Some subspecies and even species, such as E. q. danielli and Hippotigris isabellinus, were only based on illustrations (iconotypes) of aberrant quagga specimens. [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). [12] It has been pointed out that early morphological studies were erroneous; 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

Specimen in the Museum für Naturkunde, Berlin, which has been sampled for DNA

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. [8] In spite of these findings, many authors subsequently kept the plains zebra and the quagga as separate species. [5]

The mare in London Zoo, 1870

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)




Grévy's zebra (E. grevyi)





Quagga (E. q. quagga)



Damara zebra (E. q. antiquorum)- Chapman's zebra (E. q. chapmani)




Grant's zebra (E. q. boehmi)





Other Languages
Afrikaans: Kwagga
العربية: كواجا
беларуская: Квага
български: Куага
català: Quaga
čeština: Zebra kvaga
dansk: Quagga
Deutsch: Quagga
Ελληνικά: Κουάγκα
فارسی: کوآگا
Gaeilge: Cuaga
galego: Quagga
한국어: 콰가
Հայերեն: Կվագգա
Ido: Quago
Bahasa Indonesia: Quagga
עברית: קואגה
Basa Jawa: Quagga
ქართული: კვაგა
қазақша: Квагга
latviešu: Kvaga
lietuvių: Kvaga
magyar: Kvagga
മലയാളം: ക്വാഗ്ഗ
Bahasa Melayu: Quagga
Nederlands: Quagga
日本語: クアッガ
norsk: Kvagga
polski: Zebra kwagga
português: Quaga
română: Cvaga
русский: Квагга
Scots: Quagga
Seeltersk: Quagga
Simple English: Quagga
српски / srpski: Квага
srpskohrvatski / српскохрватски: Kvaga
suomi: Kvagga
svenska: Kvagga
українська: Квага
Tiếng Việt: Quagga
粵語: 擬斑馬
中文: 斑驢