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The name "Quagga" is an onomatopoeia from the sound the Quagga makes. Click the play button to hear it  

The Project

How the Project Came About, by Reinhold Rau

“Here is the Veld” by Attilio Gatti. In this book is a story of the last of the quaggas. Although the last authenticated quagga died in approximately 1883, there is an anecdotal account of a lone quagga surviving into the 1930s, related in Attilio Gatti’s lively account of adventures in Africa entitled “Here is the Veld”. However, the tale is unsupported by photographs or other confirmatory evidence or sitings and therefore must remain to some degree speculative.

These photographs, taken by Duco Quanjer of Holland, show the Artis mare that died in 1883. This exhibit is situated at the Naturalis museum in Leiden, Holland.

The origin of the name “quagga”

1quagga-paintingThis unusual name for a variety of zebra has been adopted from the Hottentot speaking indigenous people of the South African interior. The name “Quagga” has been spelt in a variety of ways, ra and Grevy Zebra (the latter which occurs only in East Africa). The name “Quagga” has been spelt in a variety of ways, according to the language in which it is used. Pronounced correctly, the double “g” as a guttural “ch”, as in the Scottish word “loch”, and with the emphasis on the first syllable.” Quagga” is an imitation of the animals call, which it shared with the other Plains Zebras. The Quagga’s nearest relative, the “true” Burchell’s Zebra, subspecies Equus quagga burchelli (also extinct), to the north of the Quagga’s distribution, became known as the “Bontquagga”. Unfortunately this distinction has often been omitted, and both forms were simply referred to as “Quagga”. Eventually the term “Quagga” became used, especially in Afrikaans, for any zebra, including the other two species, Mountain Zebra and Grevy Zebra (the latter which occurrs only in East Africa).

Extinction is forever

2quagga-extinctionIf a species of animal or plant has disappeared from the earth, either through natural causes, or through mankind’s activities, the loss is irreversible. However, the extinct Quagga was not a zebra species of its own but one of several subspecies or local forms of the Plains Zebra. Despite highly sophisticated genetic manipulations being used and reproduction in animals and plants being supported and enhanced through intricate techniques, extinction is still as final as it has always been. If a species of animal or plant has disappeared from the earth, either through natural causes, or through mankind’s activities, the loss is irreversible. However, the extinct Quagga was not a zebra species of its own but one of several subspecies or local forms of the Plains Zebra. This fact makes a big difference – the Quagga’s extinction may not be forever! An exciting breeding project has been on-going since 1987 which aims at reversing the Quagga’s extinction.

Why did the quagga become extinct?

3quagga-extinctionThe quagga’s extinction is generally attributed to the “ruthless hunting”, and even “planned extermination” by colonists. Secondly, the confusion caused by indiscriminate use of the term “Quagga”, for any zebra, prevented “last minute efforts” to save the Quagga from extinction. All members of the horse family feed predominantly on various grasses. Grasses in the Karoo and southern Free State where Quaggas occurred, are sparse. Wild grass eating animals such as the Quagga were perceived by the settlers as competitors for their sheep, goats and other livestock. Much has been written about the reasons for the extinction of the Quagga; it is generally attributed to the “ruthless hunting”, and even “planned extermination” by colonists. It’s flesh is said to have been welcome food for the farm labourers, while the skin was used as “grainbags” and “leather”. Great numbers of raw animal hides were exported during the 19th century for the leather industry. South Africa was known as a “hunters paradise”. Books such as “Portraits of the Game and Wild Animals of Southern Africa” (Harris, 1840), provide evidence of large scale killing of wild animals, done not only by the settlers, but also by those privileged to journey to the Cape of Good Hope to satisfy their lust for hunting. Such large scale hunting in South Africa during the 19th century, has drastically reduced the one time abundance of wild life, resulting in the disappearance of some species in certain areas. However, the perception that the Quagga was singled out for extermination does not seem to be supported by other historical evidence. While excessive hunting played a major role in the disappearance of the Quagga, the confusion caused by indiscriminate, that is, general use of the term “Quagga”, for any zebra, also contributed substantially. It was probably this confusion which prevented “last minute efforts” to save the Quagga from extinction. It was only realised years later that when the Quagga mare at the Artis Magistra zoo in Amsterdam died on the 12th August 1883, she was the last of her kind! The true Quagga vanished unnoticed. The indiscriminate use of the term “Quagga”, to apply to any zebra (especially in the Afrikaans language) remains an unfortunate generalisation which persists to this day.

What is a species or subspecies?

4quagga-speciesA population, however large or small, in which all individuals share basic genetic characteristics, and therefore produce fertile offspring, constitutes a species. If various populations within a huge distribution area do differ from each other in appearance, they are considered different subspecies. If a species occurs over a wide geographical area, as for example, the Plains Zebra, (north-east Africa to South Africa), populations in different parts of the distribution area, especially at the opposite ends, may look quite different from each other. Yet, when members of those populations are mated, they produce fertile offspring; that is, these offspring are able to reproduce. If various populations within a huge distribution area do differ from each other in appearance, they are considered different subspecies. If there are no geographical barriers which separate such populations or subspecies, the change in appearance is gradual and is referred to as “cline”. If however, there are geographical barriers which separate populations that were formerly part of a unified distribution, such isolated populations or subspecies could differ from others more markedly. Depending on how long they have been isolated, they may be on the verge of becoming separate species, as there is no more exchange of genes between these and other subspecies. A variety of zebra, known as the “Quagga”, inhabited the Karoo and southern Free State of South Africa well into the second half of the 19th century, when it became extinct. It differed from other zebras mainly in having been striped on the head, neck and front portion of it’s body only, and in having been brownish, rather than white, in it’s upper parts. The belly and legs were unstriped and whitish.

How was the quagga related to other zebras?

5quagga-zebraThere has never been unanimous agreement between zoologists regarding the Quagga’s relationship to other members of the horse family. Some regarded the Quagga as a full zebra species, while others treated it as the southern-most subspecies of the widely distributed Plains Zebra (often referred to as Burchell’s Zebra). While most scientists accept the Quagga as belonging to the zebras, in 1980 one researcher did suggest that the Quagga was more closely related to the horse than to the zebra. It was thought that this question about the Quagga’s relationship to other Equids, would probably never be answered, as the Quagga had long since become extinct, thus precluding the study of the living animal. Against all expectations, the question of the taxonomic status of the Quagga was answered in 1984. Three groups of scientists from the University of California undertook molecular studies on dried flesh and blood samples that had been removed from Quagga skins during re-mounting by Reinhold Rau (Taxidermist, South African Museum) of four old museum specimens in 1969/70 and 1980/81. The biochemists obtained protein and DNA fragments from the samples. The DNA fragments were successfully cloned. Both the protein and the DNA confirmed the status of the Quagga as a subspecies of the Plains Zebra. Latest (2005) Quagga DNA research results, based on small tissue samples of 13 museum specimens, confirms the subspecies status of the Quagga as obtained from tissue of one museum Quagga specimen in 1984. Read latest Quagga DNA research results, as published online by the Royal Society in “Biology Letters”, 5 July 2005.

How many different zebras are there?

6quagga-zebraThere is a lot of confusion about Burchell’s Zebra, Quagga and other zebras, despite there being only three zebra species. The reason for this is in the history of zebra descriptions and naming. Whenever an early explorer took a zebra skin from Africa to Europe, it did not match any of those in collections, so, it “needed a name”. That there is enormous individual variation in, especially, the Plains Zebra (which is often refered to as Burchell’s Zebra), had not been expected nor realized until the early 1900’s. By then, the Quagga, which had been described and named in 1788, had become extinct. The Burchell’s Zebra, described and named in 1824, was still around. Gradually, further north, somewhat more extensively-striped zebra populations became known. It was noticed that they were very similar to Burchell’s Zebra, and they were described and named as subspecies of Burchell’s Zebra. These subspecies were usually given names of explorers, like Chapman, Wahlberg, Selous, Grant, Boehm, etc. Eventually the zebra population from which William Burchell had taken a skin to the British Museum, had been wiped out, but “Burchell’s Zebra subspecies” continue to exist in many areas of Africa. Now I must explain why I prefer to speak of Plains Zebra, rather than Burchell’s Zebra, as is often done. The original Burchell’s Zebra (sometimes refered to as the “true” Burchell’s Zebra) is, or rather was, one of the subspecies of the species under discussion. Consequently, all the other subspecies (with explorers’ names) should be called Chapman’s Burchell’s Zebra, Wahlberg’s Burchell’s Zebra, Selous’s Burchell’s Zebra, and the “extinct” subspecies burchelli should be called Burchell’s Burchell’s Zebra. This would be ridiculous. Because the species that we are discussing here, lives on the plains, in contrast to the Mountain Zebra, which prefers mountainous terrain, the term “Plains Zebra” for the species as a whole, with its various subspecies (and there is no agreement among scientists how many “subspecies” there are), is a much more sensible term than Burchell’s Zebra. Fortunately this usage seems to be favoured more and more. It will certainly gradually eliminate the enormous confusion that exists. When it was realized that there are far too many names for zebras, and many were consequently made synonyms, the Quagga was no longer there. How it was related to the other zebras, was not certain. So, one left it as a species (as it had been described, after all), and called the few zebra subspecies that live on the plains, “Burchell’s Zebras”. Then there was, of course, the Mountain Zebra, and, in East Africa, the Grevy Zebra. Three living zebra species, and one extinct “species”? No one was certain about this. Some scientists tended to see the Quagga as a subspecies, others as a species. What is more, it was thought that the question about the Quagga’s taxonomic position could no longer be answered, because there were no more Quaggas around to be studied. But then, in the early 1980’s, to everybody’s surprise, that question WAS answered, through the analysing of the Quagga’s DNA from tissue that was removed during the remounting of several of the stuffed original Quaggas in museums. These developments are fairly new, and the results of the Quagga DNA analysis, namely that the Quagga WAS one of the Plains Zebra subspecies, not a species of its own, have not yet been absorbed everywhere, especially where people are not involved in Equid taxonomy. Now, was the Quagga a subspecies of Burchell’s Zebra, or the other way around? That is simple, because if it is established that two former species names in fact refer to one and the same species, then the older of the two names takes precedence over the younger. Equus quagga —1788, Equus burchelli —1824. All plains zebras therefore, including the Quagga and the “true” Burchell’s Zebra (as it is sometimes called) are subspecies of Equus quagga. The Quagga’s full name is Equus quagga quagga; its immediate northern cousin was Equus quagga burchelli; the next subspecies in a northerly direction presently is Equus quagga antiquorum, etc.

Selective breeding

7quagga-selective-breedingAmong those scientists who considered the Quagga as having been the southern-most subspecies of the Plains Zebra, were Otto Antonius, zoo director in Vienna and the two brothers, Heinz and Lutz Heck, both zoo directors, the former at Munich and the latter at Berlin, Germany. The breeding experiments of the Heck brothers, largely with domestic horses and cattle, are well known. The aim was to breed animals which resemble the wild ancestors of both the domestic horse and domestic cattle. Lutz Heck, in his 1955 book entitled “Grosswild im Etoshaland”, suggested that careful breeding with Plains Zebra, that have been selected for their brownish basic colour, and/or reduced striping, could produce an animal identical to the extinct Quagga! In 1971, Reinhold Rau visited museums in Europe to examine most of the preserved Quagga specimens, after having dismantled and re-mounted the Quagga foal at the South African Museum in Cape Town in 1969/70. During this tour he discussed the feasibility of attempting to re-breed the Quagga with Dr. Th. Haltenorth, mammalogist, at Munich, Germany. Dr. Haltenorth saw merits in such a plan and expressed his surprise that such a programme had not already been started in South Africa. Having critically examined 21 of the 23 preserved Quaggas, and being familiar with the high degree of variation in the Plains Zebra populations inhabiting the Etosha National Park in Namibia, the Kruger National Park, as well as parks in Zululand and Swaziland, Rau decided to work towards the implementation of a Quagga re-breeding programme. Contact was made in 1975 with zoologists and Park authorities, in the hope of stimulating interest in the project. Reactions to his proposals were on the whole negative, which was not surprising, considering that most English language scientific literature considered the Quagga as a separate species, a view which, if correct, would render any attempt to re-breed the Quagga a futile exercise. However, Rau did not abandon his re-breeding proposal, as he considered the Quagga to be a subspecies of the Plains Zebra. The plan received new impetus in the 1980’s by molecular studies that compared sequences of genetic code of Mitochondrial DNA extracted from tissue samples from a Quagga’s skin. Comparison of these sequences with those of the Plains Zebra, demonstrated their close affinity, at least with reference to the sequenced genes, indicating that the Quagga was a subspecies of the Plains Zebra. Then came another fortunate event. The retired veterinarian, Dr. J. F. Warning of Somerset West, contacted Rau during the latter part of 1985. He was an expert in animal husbandry and had been associated in horse and cattle breeding for more than 50 years in Germany and Namibia. He was a friend of Prof. Lutz Heck and had spent much time with him during the latter’s stay in Namibia (which resulted in Heck’s book mentioned above). Gradually a more positive attitude was taken towards the proposed Quagga re-breeding programme, as the DNA examination results appeared in publications from 1984 onward. Influential persons became involved and during March 1986 the project committee was formed. During March 1987 nine zebras, out of approximately 2 500, were selected and captured at the Etosha National Park. Their capture and arrival at the specially constructed breeding camp complex at the Nature Conservation farm “Vrolijkheid”, near Robertson, in the Cape, on 24th April 1987, marked the commencement of the Quagga re-breeding project. The first foal was born on the 9th of December 1988. During the successive years, further selected breeding stock taken from Etosha and Zululand have been added. The first foal of the second offspring generation (F2 generation) was born in February 1997. Reproductive maturity is reached only at two to three years in mares and four to five years in stallions.
The increased number of zebras led to a proportionate increase in the cost of feeding them, so much so that the limited funds of the project became stretched to the point where the breeding venue at Vrolijkheid had to be abandoned. In October 1992 six zebras were moved nearer Cape Town onto land which had sufficient natural grazing. As the new site proved to be a success, the remaining zebras from Vrolijkheid were moved there and to two additional new sites in 1993. In July 2004 Quagga Project breeding groups are living at 11 localities near Cape Town, with a total of presently 83 zebras. In addition there are 6 good stallions at 4 different places, held in reserve for replacement, should the need arise. There have been some losses, due to old age, illness or injury. Some of the less suitable offspring have been sold. It is expected that this continuous selective breeding will, with successive generations, reduce the high degree of individual variation, both in colour and in extent of striping, which are characteristics of the southern Plains Zebra. Eventually individuals should emerge whose coat-pattern characters closely resemble that of the extinct Quagga.

Criticism of the project

9quagga-museumWhile the project is progressing well, there are still those who have certain reservations or are outright against the project. The genetic basis of the Quagga Breeding Project, relies on the demonstration by Higuchi et al (1987) (Mitochondrial DNA of the Extinct Quagga: Relatedness and Extent of Postmortem Change. Journal of Molecular Evolution 25:283-287) that the mitochondrial DNA of the Quagga is identical to that of other Plains Zebras. Therefore the Quagga and other Plains Zebras belong to the same species and consequently the Quagga should be considered merely a different population (or deme), of the Plains Zebra. It has been argued that there might have been other non-morphological, genetically-coded features (such as habitat adaptations) unique to the Quagga and that therefore, any animal produced by a selective breeding programme would not be a genuine Quagga. Since there is no direct evidence for such characters and since it would be impossible now to demonstrate such characters were they to exist, this argument has limited value. The definition of the Quagga can only rest on its well-described morphological characteristics and, if an animal is obtained that possesses these characters, then it is fair to claim that it is a representation of, at least, the visible Quagga phenotype.Furthermore, since the indigenous grasses in the original habitat of the Quagga are not significantly different from those areas occupied by extant Plains Zebras, and since extant Plains Zebras occupy habitats of similar degree of aridity to those of the Quagga, there is no sound reason for proposing significant adaptive features of the Quagga to its original habitat, and no reason to believe that animals produced in the selective breeding programme would not survive successfully in the region formerly occupied by the Quagga.

Quagga material in the world’s museums

8quagga-criticismAll that remains of the countless numbers of Quaggas, that inhabited the vast plains of the Karoo and southern Free State in South Africa, are: twenty three skins, which are mounted, in more or less life-like positions, seven skeletons and some skulls and footbones, housed in Museums mostly in Europe, illustrations and descriptions made by either early travellers to Southern Africa or of Quaggas in captivity in Europe … view list of museums [ + ]

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