Quagga

Half zebra, half horse

The British naturalist William Burchell is often regarded as the discoverer of the animal that travelers described as “half zebra, half horse” or “unfinished zebra“—the quagga. However, the species had already been mentioned earlier by explorers. Even before Burchell’s “discovery” in 1812, the English naturalist George Edwards introduced the quagga to zoology in 1758 with the first volume of his Gleanings of Natural History. It contained an illustration of the animal with the caption “female zebra”. Edwards described the animal:

“This interesting animal was imported alive from the Cape of Good Hope together with a male. The male died on the way to London; I did not see it. The female, however, lived for several more years (…). The animal was unruly and grim, so that no one dared approach it.”

Vom Aussterben bedroht? 1981. p. 44. I. Akimuschkin

If Edwards had seen the male that died on the voyage to London, he would surely have noticed that it bore the same characteristic reduced stripe pattern on the hindquarters as the female.

In fact, the quagga differed in its coat coloration from modern equids such as zebras, horses, or donkeys. On the head and neck it had zebra-like brown and white stripes that usually became paler toward the torso and finally transitioned on the rear part of the body into an unstriped reddish brown reminiscent of a horse. However, this distribution of stripes varied greatly between individuals.

The Swedish naturalist Anders Sparrman spent several months in southern Africa between January and November 1772, and again in 1775, to study the flora, fauna, and the region’s ethnic groups. For the Naturhistoriska riksmuseet in Stockholm, he collected a fully developed quagga fetus, which is still preserved there today. During his time in Africa, Sparrman already realized that the quagga was a distinct form and not the female form of the zebra. In his travel report, first published in French in 1785 and in English in 1786, A Voyage to the Cape of Good Hope, he wrote:

“It was here that I saw for the first time in my life one of those animals called quaggas by the Hottentots and colonists. It is a species of wild horse, very like the zebra; the difference consisting in this, that the quagga has shorter ears, and that it has no stripes on its fore legs, loins, any of its hind parts. (…) The females of each species are marked like their respective males, excepting that the colour is somewhat more lively and definite in these latter.”

A Voyage to the Cape of Good Hope. 1786. p. 6f. A. Sparrman

William Burchell, the supposed discoverer of the quagga, mentioned the quagga in Volume 1 of his work Travels in the Interior of Southern Africa, published in two volumes in 1822 and 1824: Volume 1. His observations contributed significantly to the scientific classification of the species. He explained: “This beautiful animal has so far been mistaken by naturalists for the zebra. When these animals were first described by modern writers, the quagga was regarded as the female zebra.”

According to an article (1959) by the Dutch zoologist Adolph Cornelis van Bruggen, the exact range of the quagga was uncertain, especially regarding the northern limit. Most authors assume that the animals occurred in the semi-desert landscape of the Karoo in the Cape Province and in the southern part of the Orange Free State, today known as the Free State province, in South Africa. The Orange River likely served as the northernmost boundary—the second-longest river in southern Africa—and to the east the area was bounded by the Vaal River. It is certain that the quagga never occurred north of the Vaal. To the south, the Great Kei River or the sea probably limited the animals’ habitat.

Quagga – fact sheet

alternative names	kwagga (Afrikaans), iqwara (Xhosa)
scientific names	Equus quagga quagga, Equus burchelli quagga, Equus quagga danielli, Equus quagga lorenzi, Equus quagga greyi, Equus quagga trouessarti, Equus quagga, Hippotigris quagga, Hippotigris isabellinus
original range	South Africa
time of extinction	1883
causes of extinction	hunting, habitat loss

Why the quagga went extinct

To this day, Africa preserves the largest share of its late Pleistocene megafauna. Species such as lions, elephants, rhinoceroses, giraffes, hippos, and zebras survived, but several extinction waves of large mammals, especially at the transition from the Pleistocene to the Holocene, and European colonization of Africa led to the loss of many species. Besides the quagga, the bluebuck, the bubal hartebeest, and the Cape warthog also disappeared.

In The Doomsday Book of Animals (1981), David Day notes that quaggas were mainly distributed in the Cape Colony and the Orange Free State. This limited range made them particularly vulnerable to extinction—a fate shared by many endemic species. Farm fences further restricted the quagga’s habitat by separating groups and individual animals from one another, according to Reinhold Rau in his revised list of preserved quagga material (1974).

Meat, hides, and grain sacks

In the Cape Colony, the quagga was the only zebra in the veld—a landscape shaped by open grasslands, savannas, and partly bushy terrain. Until the 17th century, these plains were inhabited exclusively by the Khoikhoi, also known as Hottentots, an indigenous people in what is now South Africa and Namibia. With the arrival of the Boers—Dutch, Flemish, German, and French settlers — from 1652 onward, the quaggas’ fate began to change. The animals offered the settlers an obvious food source and were valuable suppliers of hides.

The Boers established huge farms, where they often forced the indigenous population to work, and hunted thousands of the wild horses. Their main interest was the hides of the quaggas, from which leather house shoes, so-called veldshoen, as well as sacks for storing grain, dried fruit, and dried meat were made. (Silverberg)

Henry Anderson Bryden, an English hunter and naturalist, reported in Kloof and Karroo (1889)—a book about the natural history of the Cape Colony—that farmers in the Orange Free State organized real hunting expeditions and safaris in the 1860s to meet the rising demand for quagga, zebra, and other wild-animal hides. Wagonloads of prepared skins were transported to the coast and sold there. The British officer and safari pioneer William Cornwallis Harris also documented in his writings how European travelers visited the Cape of Good Hope to satisfy their passion for hunting.

In The Auk, the Dodo, and the Oryx (1967), Robert Silverberg noted that Europeans did not value horse meat and therefore had little interest in using quaggas as a food source. However, quagga meat was considered a delicacy by African workers whom the settlers had to supply. To feed their workers, the Boers shot countless zebras, which thus became a staple food source for farm laborers. Even before the arrival of Europeans, the quagga had played a similar role as a staple for the Khoikhoi.

In the 1830s, the Boers moved farther north and exploited the larger populations of other zebra species there. Although these hunts were also intense, they did not lead to such a drastic decline as in the case of the quaggas.

Farms instead of habitat

Rau (1974) stated that people sometimes even spoke of a “planned extermination” of the quaggas by the colonists, as the animals were considered competition for livestock. Farmers regarded quaggas and other large herbivores such as antelopes as direct food competitors because they grazed the same grass on open pastures. Especially in the second half of the 19th century, systematic hunting expeditions drove the zebras out of farmers’ grazing areas.

With the establishment of fenced livestock pastures, the situation worsened further. The quagga’s habitat was drastically reduced, leading to fragmentation of populations. The animals could no longer move freely to find new grazing grounds or water sources. Fences also prevented encounters and reproduction between different groups, reducing genetic diversity and weakening the quaggas’ resilience to environmental change. Eloff (1966) described this problem as a “steadily impoverished gene pool” amid further migration to the south and decreasing adaptability to changing environmental conditions.

The drought years after 1876 further accelerated the quaggas’ decline. Rau (1974) refers to Alfred de Jager Jackson, who described the dramatic consequences of this period in his book Manna in the Desert (1920). Jackson, who spent his youth on a farm in South Africa, wrote: “As I said, the good years ended in 1876. The following year was a time of terrible disease and drought. The carcasses of thousands of dead animals rotted across the land.” This combination of overhunting, habitat loss, and extreme climatic conditions ultimately led to the quagga’s final disappearance.

Taming and human use

Quaggas were hunted not only for sport and as a source of meat or as suppliers of hides and skins, but also for other purposes. As David Day reports, colonists used the animals as “guard dogs” for their livestock herds. Quaggas were supposed to deter attackers such as hyenas or wild dogs by aggressively defending themselves against intruders and raising the alarm.

Despite their nature often described as stubborn and aggressive, people succeeded in taming quaggas in individual cases. One example is Sheriff Parkins, who in the 1830s in London harnessed two quagga stallions as draft animals in front of his carriage.

When did the quagga disappear?

The quagga’s extinction initially went largely unnoticed by zoologists and conservationists. This was also due to linguistic confusion: in Afrikaans, the term “Kwagga” referred not only to the true quagga but also to plains and mountain zebras. Only outside the Cape Colony, as Reinhold Rau (1974) describes it, was the name “Quagga” in its various spellings applied specifically to the true quagga.

The last known quagga lived in the Artis Zoo in Amsterdam from May 9, 1867, and died there on August 12, 1883. At that time, no one suspected that it was the last specimen of its kind. The zoo even requested another animal, assuming that quaggas might still live in remote regions of the Cape Colony. In Stripes Faded, Barking Silenced: Remembering Quagga (2014), Rick De Vos writes that the quagga’s extinction was officially recognized only in 1900 through the London Convention for the Protection of Wild Animals, Birds, and Fish in Africa.

In the 16th century and at the beginning of the 17th century, there are said to have been large herds of quaggas. The Soviet zoologist Igor Akimushkin vividly describes this abundance:

“Back then, there were supposedly so many quaggas at first that the Boers ran short of lead during hunting. So they cut the bullets out of the killed animals, reloaded their weapons with them, and fired at the next defenseless animal, which had not even fled far.”

Vom Aussterben bedroht? 1981. p. 47. I. Akimuschkin

But as early as the middle of the 19th century, a different trend became apparent. The British officer Sir William Cornwallis Harris, who stayed in South Africa between March 1836 and December 1837, wrote in his 1840 book Portraits of the Game Animals of Southern Africa:

“[The quagga] was formerly extremely common within the Colony, but vanishing before the strides of civilization, is now to be found in very limited numbers, and on the borders only.”

The Auk, the Dodo, and the Oryx. Vanished and Vanishing Creatures. 1967. p. 104. R. Silverberg

Harris noted that populations in the north of the colony were more stable because hunting there was less intensive. Nevertheless, increasing hunting began there as well. According to Silverberg, by 1850 the Boers had shot most of the wild animals in their settlement area, including the quaggas. After that, they moved farther north to hunt still untouched herds. This continued for another 20 years until the last animal in the wild, according to Silverberg, was killed around 1870.

According to Akimushkin, the last two quaggas of the Cape Province were shot in 1850 in the Tygerberg hills in western South Africa. The last wild population probably persisted until 1878 in a remote semi-desert region in the Orange Free State. After the devastating drought from 1777 onward, the quagga was presumably extinct in the wild.

The exact timing of extinction is given differently in the literature. The German zoologist Alfred Brehm wrote in Brehms Thierleben (1911) that the quaggas in the Orange Free State became extinct in 1870 and that the species was completely eradicated by 1879. Henry Anderson Bryden, a naturalist and writer, dated the extermination in the Cape Colony to 1865 to 1870, and in the Orange Free State — the animals’ last refuge — to 1870 to 1873.

Quagga sightings in the 20th century

As with many extinct animal species, there were also repeated alleged sightings of the quagga after its disappearance. For some time, these reports fueled speculation about the species’ possible survival. Victor Franke, the last commander of the Imperial Schutztruppe in German South West Africa (present-day Namibia), noted in his diaries that he claimed to have seen small herds of quaggas in 1901. Despite his account, no proof of this observation was ever found.

In 1917, Major C. N. Manning, a British officer, reported an encounter with a herd of unusual animals in the Kaokoveld, a remote region in present-day Namibia. These animals, according to Manning, resembled zebras in size and shape but had a brown coloring and stripes limited to the front part of the body. Without binoculars, he could not make out further details.

Manning reported his sighting to the local government official C. M. Hahn, who then initiated a search for the mysterious animals. However, the efforts were unsuccessful. Hahn concluded that Manning had probably seen Hartmann’s mountain zebras (Equus zebra hartmannae). Hahn explained:

“In the hot season, generally at midday, with the sun is overhead, Hartmann’s zebras from a distance resemble dark brown donkeys. When the heat haze is severe, it is difficult, at times impossible, to discern the dark markings without the aid of binoculars.”

The Auk, the Dodo, and the Oryx. Vanished and Vanishing Creatures. 1967. p. 107. R. Silverberg

In the years after Manning, explorers, hunters, and locals also reported sightings of brown wild horses with reduced stripe patterns. According to Silverberg, the most recent alleged quagga sighting was reported in 1940.

Besides the theory that the animals seen were Hartmann’s mountain zebras, Akimushkin suggests that observers may have seen zebroids—hybrids between donkeys and zebras—which occasionally occur in the wild in Africa when animals live in close proximity. Such hybrids often have a striking, irregular stripe pattern that recalls the quagga.

Brehm also points out that the linguistic confusion—in which the term “quagga” was traditionally used in South Africa for all zebras—likewise contributed to the ongoing reports of alleged sightings. Brehm remarked: “Continued claims that it still lived had no significance.”

Breeding back: bringing the quagga back to life?

The Quagga Project was founded in 1987 by Reinhold Rau. The aim of the project is to recreate the extinct quagga through selective back-breeding. Rau cataloged preserved museum specimens and, in the 1980s, collected skin samples that enabled DNA analyses. These confirmed that the quagga is closely related to the plains zebra and was probably a subspecies of the plains zebra.

Inspired by the work of Heinz and Lutz Heck in the 1920s—who, among other things, dealt with the back-breeding of extinct aurochs—Rau began a similar program to revive the quagga: Through the targeted selection of plains zebras with reduced stripes, animals are to be produced over several generations that resemble the original quagga in appearance. Instead of cloning the species, the scientists relied on selective breeding to recreate the typical brown-and-white pattern and the missing stripes.

The resulting “Rau quaggas”, which do indeed show a reduction in striping, are genetically closely related to southern plains zebras. By the fifth generation, there were already about 90 animals that looked very similar to the extinct quagga. In 2022, Süddeutsche Zeitung reported that the population is now around 200 animals. More recent official figures are not available. However, it can be assumed that the population has continued to grow since then, as the project is ongoing.

Part of the project is also the reintroduction of the bred Rau quaggas into the Karoo in South Africa. While some researchers see the animals as an authentic quagga reconstruction, others criticize them as simply zebras with fewer stripes that look like the original. Essentially, Rau quaggas are an imitation; the genetic traits of the true quagga are irretrievably lost. One scientific benefit of the project could be developing a method to recreate populations of threatened species in order to refill the ecological gap left by extinct species or populations.

Evolution and taxonomy of the quagga

The name “quagga” derives from the language of the Khoikhoi, an indigenous population group in southern Africa. Originally, the “gg” in the name was pronounced like a “ch”, supposedly imitating the animal’s characteristic sound. In his Gleanings of Natural History (1758), George Edwards described this sound as follows: “The animal’s cry bore no resemblance to that of a donkey; it rather reminded one of the barking of a mastiff.” Since no audio recordings of the quagga exist, its sounds can hardly be reconstructed today. Whether the name is indeed an onomatopoeic imitation of its call therefore cannot be conclusively clarified.

Early classifications: species or subspecies?

The original scientific description of the quagga was published in 1778 by the Dutch naturalist Pieter Boddaert; he classified the quagga, Equus quagga, as an independent species. Traditionally, the quagga as well as plains and mountain zebras were placed in the subgenus Hippotigris. The precise taxonomic position of the quagga fueled debates for decades.

The British zoologist Reginald Innes Pocock proposed in 1904, in The Cape Colony Quaggas, to regard the quagga as a subspecies of the plains zebra (E. quagga, previously E. burchelli). He doubted that it represented an independent or a fourth zebra species. Although there are some clear physical differences between the quagga and other plains zebras, these differences were not strong enough to justify a separate species. He also argued that observed morphological variation in specimens—such as stripe patterns and coloration—falls within the range of variation found in other plains zebra subspecies. The quagga thus received the scientific designation E. q. quagga.

Variability in coat pattern

Unlike zebras with light bodies and dark stripes, the quagga was a brownish animal with a limited stripe pattern that mainly affected the front part of the body. The rear parts of the body were usually completely stripe-free.

The species showed high variability in stripe patterns—some animals had almost no stripes, while in others most of the body, except for the hind parts, legs, and belly, was striped. For example, in the Amsterdam museum specimen the stripes do not extend beyond the shoulder, whereas in the Leiden specimen they reach the rear part of the body. And the mounted skin in Milan has only a few light stripes in the neck area, so that at first glance it resembles a donkey. This variability in coat pattern led to several subspecies being described based on the extent of the stripe pattern.

Bruggen wrote that the few preserved specimens would not provide enough material to justify a subdivision into subspecies. He also notes that differences in zebra coat patterns are purely phenotypic. Some alleged subspecies, such as E. q. danielli, were based solely on historical illustrations of individual extraordinary specimens.

Morphological studies

There are only a few fossils of the quagga—or rather, assigning fossil remains is difficult, because at the time the bones were collected, the designation “quagga” was historically used for all zebras. Morphological studies of skulls and other skeletal parts produced contradictory results.

In a study published in 1991, the paleontologist Augusto Azzaroli and the evolutionary biologist Roscoe Stanyon concluded from morphological examinations that the quagga and the plains zebra were not only different species but also not closely related. Another analysis of skull measurements from 1999 likewise suggested that the quagga differed from the plains zebra as much as the plains zebra differed from the mountain zebra (E. zebra). However, the explanatory power of this study is limited, as the researchers Richard G. Klein and Kathryn Cruz-Uribe had only four skulls and specimens of questionable origin available.

In 2004, the Australian anthropologists Colin P. Groves and Catherine Hobart Bell revised the taxonomy of zebras by applying traditional taxonomic methods such as skull measurements and comparing coat patterns. Like Pocock before them, they concluded that the quagga was not an independent species but rather a subspecies of the plains zebra. They grouped Burchell’s zebras and plains zebras under the designation Equus quagga, because no absolute differences between the groups could be determined. In their reassessment, they identified six subspecies of the plains zebra, one of which is the quagga.

Genetic studies

The quagga was the first extinct animal whose DNA was analyzed. In 1984, the geneticist Russel Higuchi and his colleagues wanted to find out whether DNA is preserved in the remains of extinct animals and can be extracted from them. For the study, they examined a museum specimen from which they were able to take a small amount of DNA from dried muscle tissue. The researchers compared the animal’s mtDNA sequences with those of other equids and found differences from the mtDNA of the mountain zebra, suggesting that quagga and mountain zebra split from a common ancestor about three to four million years ago. From this, the researchers concluded that the quagga is more closely related to zebras than to horses.

A year later, an immunological study (1985) by the biochemist John M. Lowenstein showed that proteins preserved in museum specimens are much more similar to the serum proteins of the plains zebra than to those of other modern zebra species. This analysis also suggests that the quagga is closest to the plains zebra.

A study by Jennifer A. Leonard (2005) complemented and expanded the results of Higuchi’s original work by conducting a genetic analysis based on multiple quagga specimens. While Higuchi in the 1980s could extract only a small amount of mtDNA from a single specimen and compare it with other equids, Leonard’s team analyzed DNA from eight quaggas and one plains zebra and obtained more detailed results on genetic relatedness.

The results suggested that the quagga split from the plains zebra about 120,000 to 290,000 years ago, possibly as an adaptation to climatic changes in the Pleistocene. This supported the hypothesis that it was more likely a subspecies of the plains zebra than an independent species.

Neither species nor subspecies—a regional variation

A genetic study on plains zebra populations from 2018 once again confirmed that the quagga belongs to the plains zebra species. However, this investigation suggests that the groups previously described as plains zebra subspecies may be geographically shaped populations rather than clearly defined subspecies. The Danish biologist Casper-Emil T. Pedersen and his team found no evidence that the southern plains zebra populations—including the quagga—show enough genetic differences to be regarded as separate subspecies. Instead, the various “subspecies” of the plains zebra seem to be variations within a single species that arose through adaptation to different habitats.

The quagga appears to be genetically closer to the neighboring southern zebras of Namibia than to northern populations in Uganda. These results support the assumption that today’s plains zebras originated in southern Africa and spread about 370,000 years ago.

Current taxonomic classification

Most authors today take the view that the quagga is not an independent species but a subspecies of the plains zebra. Nevertheless, more recent genetic research suggests that the traditional plains zebra subspecies—including the quagga—are better understood as geographically shaped populations within a single species. In that sense, the quagga would largely be regarded as a regionally adapted variant of the plains zebra, which adapted its coat color and stripe reduction to the specific living conditions in southern Africa.

Why the quagga lost its stripes

Over time, scientists repeatedly asked why, unlike other zebras, the quagga had no stripes on its hindquarters. Several hypotheses have been established that are also meant to explain coat patterns in other zebra forms — such as climatic and geographic adaptation for protection from predators, defense against biting flies, or thermoregulation. Various studies have investigated why zebras show stripe patterns of different intensity and how environmental factors influence stripe variation in plains zebras.

Stripes for thermoregulation

The 2015 study How the Zebra Got its Stripes concluded, roughly, that temperature has a significant influence on stripe formation, whereas the hypotheses about camouflage from predators or protection from biting flies were less convincing. If zebras did in fact evolve their stripes for cooling, the quagga’s reduced stripes could be explained by living in cooler regions where fewer stripes were needed for thermoregulation. A counterargument, however, is that the mountain zebra also lives in cooler regions and is nevertheless strongly striped.

Fewer biting flies = fewer stripes?

Did the quagga have fewer stripes because there were fewer biting flies to deter? A study by Tim Caro (2019) showed that zebras are well protected by their stripes against parasitic flies such as tsetse flies—but only in combination with their active behavior. Stripes do not prevent flies from approaching from a distance, but they make landing nearby more difficult. In addition, zebras actively drive flies away through frequent tail swishing and running off. This combination of stripes and active behavior ensures that only very few flies can suck blood from the zebra. Since the quagga lived in regions with fewer flies, selection pressure for stripe formation may have been lower, which could explain the loss of stripes on the rear part of the body.

Fewer stripes due to geographic isolation and climatic adaptation

Leonard’s study (2005) appears the most plausible for explaining why the quagga had fewer stripes. It suggests that the quagga’s appearance—reduced stripes in the rear area and a brownish color—could have evolved relatively quickly in a dry habitat. A combination of geographic isolation and climatic adaptation could explain this rapid change in coat pattern. The researchers suspect that the loss of stripes was reinforced by the isolation of the quagga population from other zebra populations.

Color variability could be due to adaptation to drier, more open habitats during the Pleistocene, which led to a reduction of stripes. In open, drier regions of southern Africa, less conspicuous coat patterns could have been advantageous. These areas offered little cover, and a less striped, brownish coat could have camouflaged the quagga better in this environment. This evolution may have been reinforced by the last major climate changes, which interrupted patterns of gene flow. Plains zebra subspecies tend to show fewer stripes the farther south they live, and the quagga was the southernmost of these subspecies and represented the extreme of this development.

Quaggas in European zoos

The quaggas kept in European zoos in the 18th and 19th centuries were initially hardly perceived as threatened, because the term “quagga” was broadly used at the time and often applied to other striped equids as well . Only years after the death of the quagga mare in the Amsterdam Zoo in 1883 did people realize that she had been the last known specimen of her kind. By then it was too late for protective measures or breeding attempts. Had the threat been recognized in time, one might have tried to protect quaggas in South Africa or to establish captive breeding from European stock.

According to Silverberg and Akimushkin, at least 16 quaggas were shipped to Europe during the 18th and 19th centuries, beginning with the female that George Edwards described in 1758. At first they were mainly curiosities for wealthy private individuals. Only later did some of the animals end up in zoological gardens.

Kept in Paris, Berlin, and London

According to Zootierliste, holdings in Germany, Belgium, the Netherlands, France, and England are documented for European zoos. One of the earliest zoos to keep a quagga was the Jardin des Plantes in Paris. A 16-year-old stallion lived there from 1794 to 1798. Because of its advanced age, it died only a few years after its arrival. The Berlin Zoo housed two quaggas, one from 1863 to 1867 and another from 1872 to 1877—the latter may have been the penultimate animal of its kind.

In the 19th century, three quaggas lived in the London Zoo at different times, according to Silverberg. Particularly noteworthy is a pair kept by the Zoological Society of London between 1851 and 1858. The female quagga was acquired in 1851, and in 1858 Sir George Grey presented the Society with a male specimen. However, the hope of establishing a breeding program with these animals was not fulfilled.

The breeding stallion died in 1864 after, in a fit of rage, ramming into the wall of his enclosure. The mare lived until 1872 and for a long time was considered the only quagga ever photographed alive. Five photographs from 1863 to 1870 document her existence. After her death, her skeleton was given to the Peabody Museum; her skin is now in the National Museum of Scotland in Edinburgh.

Further holdings and breeding attempts

Quaggas were also kept in other European zoos and menageries. At the zoo in Antwerp, Belgium, two quaggas were reportedly even born—possibly the only documented captive births.

In Stuttgart, in the Royal Zoo (today Wilhelma), at least one quagga lived, imported in 1812. The zoo closed in 1817, and further details about this animal’s fate are not recorded. The Dresden Zoo may also have been among the institutions that kept a quagga.

In the menagerie of the Earl of Derby in London, a single animal was kept from 1850 to at least 1861. It was crossed with a Mongolian wild ass (Equus hemionus hemionus)—one of the few documented interspecific crosses.

The last known quagga lived in the Amsterdam Zoo Natura Artis Magistra from 1867 onward. The mare died on August 12, 1883, without her significance being recognized at first.

Photos of living quaggas—a rarity

Until recently, the five photos taken by Frank Haes and Frederick York between 1863 and 1870 of a quagga mare in the London Zoo were considered the only known images of a living animal of this species. In an article published in 2024, the biologist Branden Holmes and the artist Jasper Hulshoff Pol report the rediscovery of another quagga photo, taken on April 8, 1864, by the German physician Gustav Theodor Fritsch in South Africa. This image, a so-called stereophoto with two slightly offset perspectives, was listed in an old auction catalog from 1879. Hulshoff Pol came across this reference in December 2022 and was able to acquire a copy of the photo.

The newly found image shows a quagga on the “Quaggafontein” farm and provides a rare depiction of the animal in its homeland. It differs clearly from the London images: the specimen in Fritsch’s photo shows strongly reduced stripes, an important trait that distinguishes it from Burchell’s zebra. Interestingly, on the reverse of the image the animal is incorrectly labeled “bonte Quagga” (Burchell’s zebra). Peter Heywood, an expert on the quagga, confirmed that the photo shows the true quagga and not Burchell’s zebra as the label suggests, “because stripes on the legs are missing and the stripes on the body are reduced”. The incorrect label may have contributed to the significance of this photo remaining unnoticed for a long time.

Because Fritsch’s stereophoto produced two images from slightly different angles, and because of the diapositiv image of the stereophoto, the total number of known images of living quaggas increases, according to Holmes, to at least eight or nine: five shots from London, including at least one stereophoto shot (making at least six images), one stereophoto (two images), and one photoxylograph (one image). If the stereophoto was the basis for the photoxylograph, then there are eight quagga photos.

Quaggas in museums and collections

Worldwide, 23 mounted quagga skins, at least seven complete skeletons and 13 skulls, as well as other remains of the extinct animal, are preserved. They are held in museums and collections in Europe, Africa, and North America. Ten mounted skins alone are kept in German museums. However, over the course of history, some specimens have been irretrievably lost, especially during World War II. According to Reinhold Rau (1978), a mounted quagga in a museum in Königsberg (today Kaliningrad, Russia) was destroyed by a fire at the end of the war. In addition, two quagga skulls were lost during air raids on London.

The preserved specimens are not only museum exhibits, but also of great scientific importance. Modern genetic and morphological analyses of these remains have helped to better understand the quagga’s evolution and clarify its relationship to today’s zebra species. Genomic studies in particular made it possible to investigate the genetic proximity between the quagga and the plains zebra. These insights inspired Reinhold Rau in his attempt to “reconstruct” the species through selective breeding.

Despite the documented specimens, it is possible that undiscovered quagga preparations still exist in museums or private collections. However, a 1901 article titled Some Animals Exterminated During the Nineteenth Century pointed out that quaggas were not particularly attractive trophies for hunters. This makes it unlikely that there are significant additional specimens in private collections.

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