The quagga (Equus quagga quagga) was a subspecies of the plains zebra that lived only in South Africa. It became extinct in the late 1800s because people hunted it. Scientists once thought it was a separate species, but DNA studies showed it was a type of plains zebra. A later study suggested it was the southernmost group of plains zebras.

The quagga was about 257 cm (8 ft 5 in) long and 125–135 cm (4 ft 1 in–4 ft 5 in) tall at the shoulders. It had fewer stripes than other zebras, mostly brown and white on the front part of its body. Its back was brown with no stripes and looked more like a horse. The number and placement of stripes varied between individuals. Little is known about its behavior, but it may have lived in groups of 30–50. Quaggas were described as wild and lively but calmer than Burchell's zebra. They once lived in large numbers in the Karoo region of the Cape Province and the southern part of the Orange Free State in South Africa.

After Europeans settled in South Africa, quaggas were hunted heavily because they competed with farm animals for food. Some were taken to European zoos, but breeding efforts failed. The last wild quaggas lived in the Orange Free State, and they disappeared from the wild by 1878. The last quagga in captivity died in Amsterdam on August 12, 1883. Only two living quaggas were ever photographed, and only 23 skins remain today. In 1984, the quagga became the first extinct animal to have its DNA studied. The Quagga Project has tried to breed Burchell's zebras with similar stripe patterns to the quagga.

Taxonomy

The name "quagga" is believed to come from a Khoikhoi word for zebra, which sounds similar to the quagga's call, often written as "kwa-ha-ha," "kwahaah," or "oug-ga." This name is still used today for the plains zebra.

The quagga was first classified as a separate species, Equus quagga, in 1778 by Dutch naturalist Pieter Boddaert. Traditionally, the quagga and other plains and mountain zebras were grouped in the subgenus Hippotigris. Scientists have long debated whether the quagga was a separate species or a subspecies of the plains zebra. In 1902, British zoologist Reginald Innes Pocock suggested the quagga was a subspecies of the plains zebra. Because the quagga was named first, its scientific name is E. quagga quagga, while other plains zebra subspecies are called E. quagga.

Taxonomy of the quagga became more complicated because the extinct southern population of Burchell's zebra (Equus quagga burchellii) was once thought to be a separate subspecies or even a full species (E. burchellii). The northern population, known as the "Damara zebra," was later named Equus quagga antiquorum. Later, scientists realized these were the same group, so the Damara zebra is now called E. q. burchellii. The extinct southern population was once thought closely related to the quagga because both had limited striping on their hindquarters. In 1934, Shortridge placed both in the now-disused subgenus Quagga. Most experts now believe these two subspecies represent opposite ends of a gradual change in traits.

Early researchers recognized different plains zebra subspecies, but confusion existed about which were valid. Differences in striping patterns were once used to describe subspecies, but these differences were later found to be variations within the same population. Some subspecies, like E. q. danielli and Hippotigris isabellinus, were based only on illustrations of unusual quagga specimens. A 1980 study of skulls suggested the quagga was related to horses, but this was later found to be incorrect. Studying skulls from stuffed specimens can be tricky because early taxidermists sometimes used donkey or horse skulls when originals were unavailable.

The quagga has few fossils, and their identification is unclear because the name "quagga" once referred to all zebras. Fossil skulls of Equus mauritanicus from Algeria were thought to be related to the quagga and plains zebra, but they are too damaged for certain conclusions.

The quagga was the first extinct animal to have its DNA studied in 1984, which started the field of ancient DNA analysis. This study showed the quagga was more closely related to zebras than horses, sharing an ancestor with the mountain zebra 3–4 million years ago. A 1985 study confirmed the quagga was closest to the plains zebra. A 1987 study found the quagga's mtDNA diverged about 2% per million years, similar to other mammals, and again showed its close relation to the plains zebra.

Later studies had different results. A 1999 analysis of skull measurements suggested the quagga was as different from the plains zebra as the plains zebra is from the mountain zebra. A 2004 study of skins and skulls found the quagga was a subspecies of the plains zebra, not a separate species. Despite this, many scientists continued to classify the quagga and plains zebra as separate species.

A 2005 genetic study confirmed the quagga was a subspecies of the plains zebra. It showed the quagga had little genetic diversity and diverged from other plains zebra subspecies 120,000 to 290,000 years ago, during the Pleistocene. Its unique coat pattern may have evolved quickly due to isolation or adaptation to drier environments. Plains zebra subspecies have less striping the farther south they live, and the quagga lived the farthest south. Other African animals also split into species or subspecies during this time due to climate changes.

A simplified diagram based on the 2005 study shows the relationships between zebra species:

• 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)

A 2018 study of plains zebra populations confirmed the quagga was part of that species. It found no evidence of subspecies based on differences in southern populations, including the quagga. Modern plains zebra populations may have originated in southern Africa, and the quagga is less different from nearby populations than the northernmost group in Uganda. The study showed a genetic pattern from north to south, with Ugandan zebras being the most distinct. Zebras from Namibia are genetically closest to the quagga.

Description

The quagga was believed to be about 257 cm (8 ft 5 in) long and 125–135 cm (4 ft 1 in – 4 ft 5 in) tall at the shoulders. Measurements from skins showed that female quaggas were much longer and slightly taller than males, while male zebras in living populations are the largest. Its coat pattern was different from other horses: it looked like a zebra on the front but more like a horse on the back. It had brown and white stripes on the head and neck, brown fur on the upper body, and a white belly, tail, and legs. The stripes were most visible on the head and neck and became less clear toward the body, blending with the reddish brown of the back and sides until disappearing along the spine. It also had a wide dark stripe running along its back and a standing mane with brown and white stripes.

The quagga showed a wide variety of appearances, with some having very few stripes and others having patterns similar to the extinct southern group of Burchell's zebra, where stripes covered most of the body except the hind parts, legs, and belly. Based on photos and written descriptions, many people believe the quagga’s stripes were light on a dark background, unlike other zebras. Reinhold Rau, a German naturalist who started the Quagga Project, said this was an illusion, claiming the base color was creamy white and the stripes were thick and dark.

Living at the southern edge of the plains zebra’s range, the quagga had a thick winter coat that shed each year. Its skull had a straight shape and a curved space between the teeth, and it was broad with a narrow back part of the skull. Like plains zebras, the quagga did not have a dewlap (a fold of skin on the neck) like mountain zebras. A 2004 study found that the skeletal features of the southern Burchell’s zebra population and the quagga were very similar, making them impossible to tell apart. Some specimens had patterns between the two, and today’s Burchell’s zebra population still shows limited striping. This means the two groups gradually changed into each other over time. Today, some stuffed quagga and southern Burchell’s zebra specimens are so similar that they cannot be clearly identified, as no location information was recorded for them.

Behaviour and ecology

The quagga was a type of plains zebra that lived farthest south, mainly in areas south of the Orange River. It was a grazer, meaning it ate grass, and its habitat was limited to grasslands and dry scrub areas in the Karoo region of South Africa. Today, these areas are part of the provinces of Northern Cape, Eastern Cape, Western Cape, and the Free State. These regions had unique plants and animals, with many species found only there. Quaggas often gathered in groups of 30–50 and sometimes moved in straight lines. They may have lived in the same area as Burchell’s zebra between the Vaal and Orange rivers, but there is no proof they interbred. They might also have shared a small part of their habitat with Hartmann’s mountain zebra (Equus zebra hartmannae).

Little is known about quagga behavior in the wild, and old reports sometimes refer to unclear zebra species. The only clear description of quaggas in the Free State comes from William Cornwallis Harris, a British engineer and hunter. His 1840 account states:

The quagga did not live north of the Vaal River. It was once very common in the area but became rare due to human activity. Now, it is found only in small numbers near the edges of its former range. In the hot, wild plains where few people live, quaggas moved in large groups. They did not mix with other zebras but often traveled with white-tailed gnus and ostriches, especially enjoying the company of ostriches. Moving slowly across the horizon, quaggas made loud, barking sounds, and their long lines reminded travelers of caravans on the move. Large groups often traveled from dry, empty plains to find greener areas during summer, where plants grew lush and colorful.

The purpose of zebra striping has been debated, and it is unclear why quaggas lacked stripes on their hind legs. Some theories suggest stripes help hide zebras from predators, reduce fly bites, or aid in social interactions. Differences in hindquarter stripes may have helped zebras recognize their own kind during group movements. Another idea is that striping helps zebras cool down, but quaggas may have lost their stripes because they lived in cooler areas. However, mountain zebras, which also live in cool regions, have bold stripes, making this theory less certain. A 2014 study supported the idea that stripes help avoid flies, and quaggas lived in areas with fewer flies than other zebras.

A 2020 study suggested that quagga females were larger than males, possibly due to cold and dry conditions in the Karoo plateau. These conditions were even harsher in the past, such as during ice ages. Cold and dry environments may have influenced quagga evolution, including their coat color and size differences. Since female zebras are often pregnant or nursing, being larger could help them survive when food was scarce. Differences in size and coat color may also have developed due to genetic changes from isolation, but these factors may have worked together.

Relationship with humans

Quaggas have been found in cave art made by the San people, who lived in Southern Africa long ago. These animals were easy to find and hunt, so early Dutch settlers and later Afrikaners hunted them for meat and their skins. The skins were sold or used for profit. Quaggas were at risk of disappearing because they lived in a small area. Farmers used them to protect their animals, as quaggas would attack people who tried to steal livestock. Quaggas were described as energetic and nervous, especially male quaggas. They were kept in European zoos, and people tried to breed them in captivity. This effort stopped when a male quagga died by hitting its head against a wall after becoming angry. In zoos, quaggas were calmer than Burchell's zebras. One quagga lived in captivity for 21 years and 4 months before dying in 1872.

Quaggas were considered good candidates for domestication because they were calm and strong. Dutch settlers in South Africa thought this might help, as their horses struggled with the local climate and a disease called African horse sickness. In 1843, a naturalist named Charles Hamilton Smith wrote that quaggas were best suited for domestication because of their strength and calm nature. Some reports mention quaggas being tamed in South Africa. In Europe, two male quaggas were used to pull a carriage in London during the early 1800s.

To try domesticating quaggas, a British lord named George Douglas, 16th Earl of Morton, bred a male quagga with a female horse that had some Arabian blood. This produced a female with stripes on her back and legs. The horse was later sold and bred with another horse, creating offspring with zebra-like stripes. This experiment was described in 1820 by the Royal Society. It is not known what happened to the first hybrid. This led to ideas about how traits are passed down, which Charles Darwin later called pangenesis. In the late 1800s, a scientist named James Cossar Ewart disagreed with these ideas and showed through experiments that zebra stripes could appear suddenly in offspring.

There are 23 known preserved quagga specimens worldwide, including a baby, two young ones, and a fetus. Other items kept include a head and neck, a foot, seven full skeletons, and tissue samples. A 24th specimen was destroyed during World War II in Germany, and many bones have been lost. A quagga at London Zoo was once thought to be the only one photographed alive. Five photos of this quagga exist, taken between 1863 and 1870. In 2022, two old photos of a quagga were found. They were taken in 1864 by a German scientist named Gustav Theodor Fritsch. He used them to make a drawing in his 1868 book. Some people questioned whether the photo showed a quagga or a Burchell's zebra, but scientists later confirmed it was a quagga.

Quaggas disappeared from most of their habitat by the 1850s. The last wild quaggas in the Orange Free State were lost by the late 1870s. The last wild quagga died in 1878. The quagga in London died in 1872, and the one in Berlin died in 1875. The last quagga in captivity, a female in Amsterdam, lived from 1867 to 1883. Her death was not recognized as the end of the species at the time, and the zoo asked for more quaggas. People believed they might still live in the interior of South Africa. Because locals used the word "quagga" for all zebras, this may have caused confusion.

The extinction of the quagga was officially recognized in the 1900 Convention for the Preservation of Wild Animals. A Dutch stamp from 1988 featured the last quagga. The specimen is preserved in a museum in Leiden, Netherlands, and is sometimes displayed. In 1889, a naturalist named Henry Bryden wrote that the quagga’s disappearance was a shame for modern society.

After scientists found that quaggas are closely related to plains zebras, a project called the Quagga Project began in 1987 in South Africa. The goal was to breed plains zebras to look like quaggas by reducing their stripes. These zebras are called "Rau quaggas" to distinguish them from wild zebras. The project started with 19 zebras from Namibia and South Africa, chosen because they had fewer stripes on their bodies and legs. The first baby quagga-like zebra was born in 1988. If enough quagga-like zebras are created, they may be released into the Western Cape.

Introducing these zebras could help restore ecosystems by working with other animals like wildebeest and ostriches. These animals historically lived together and helped maintain grasslands. In 2006, third- and fourth-generation quagga-like zebras looked similar to old photos and preserved quaggas. This method of breeding is called "breeding back." It is controversial because the zebras will look like quaggas but will not be genetically the same. Scientists have not yet developed the technology to use DNA from extinct animals to clone them.