Zebras (, ) (subgenus Hippotigris) are African equines with distinctive black-and-white striped coats. There are three living species: Grévy's zebra (Equus grevyi), the plains zebra (E. quagga), and the mountain zebra (E. zebra). Zebras share the genus Equus with horses and asses, the three groups being the only living members of the family Equidae. Zebra stripes come in different patterns, unique to each individual. Several theories have been proposed for the function of these patterns, with most evidence supporting them as a deterrent for biting flies. Zebras inhabit eastern and southern Africa and can be found in a variety of habitats such as savannahs, grasslands, woodlands, shrublands, and mountainous areas.
Zebras are primarily grazers and can subsist on lower-quality vegetation. They are preyed on mainly by lions, and typically flee when threatened but also bite and kick. Zebra species differ in social behaviour, with plains and mountain zebra living in stable harems consisting of an adult male or stallion, several adult females or mares, and their young or foals; while Grévy's zebra live alone or in loosely associated herds. In harem-holding species, adult females mate only with their harem stallion, while male Grévy's zebras establish territories which attract females and the species is promiscuous. Zebras communicate with various vocalisations, body postures and facial expressions. Social grooming strengthens social bonds in plains and mountain zebras.
Zebras' distinctive stripes make them among the most recognisable mammals. They have been featured in art and stories in Africa and beyond. Historically, they have been highly sought by exotic animal collectors, but unlike horses and donkeys, zebras have never been completely domesticated. The International Union for Conservation of Nature (IUCN) lists Grévy's zebra as endangered, the mountain zebra as vulnerable and the plains zebra as near-threatened. The quagga (E. quagga quagga), a type of plains zebra, was driven to extinction in the 19th century. Nevertheless, zebras can be found in numerous protected areas.
Etymology The English name "zebra" derives from Italian, Spanish or Portuguese. Its origins may lie in the Latin equiferus, meaning "wild horse". Equiferus appears to have entered into Portuguese as ezebro or zebro, which was originally used for a mysterious equine reported in the wilds of the Iberian Peninsula during the Middle Ages. In 1591, Italian explorer Filippo Pigafetta recorded "zebra" being used to refer to the African animals by Portuguese visitors to the continent. In ancient times, the zebra was called hippotigris ("horse tiger") by the Greeks and Romans.
The word zebra was traditionally pronounced with a long initial vowel, but over the course of the 20th century the pronunciation with the short initial vowel became the norm in British English. The pronunciation with a long initial vowel remains standard in American English.
Taxonomy
Zebras are classified in the genus Equus (known as equines) along with horses and asses. These three groups are the only living members of the family Equidae. The plains zebra and mountain zebra were traditionally placed in the subgenus Hippotigris (C. H. Smith, 1841) in contrast to the Grévy's zebra which was considered the sole species of subgenus Dolichohippus (Heller, 1912). Groves and Bell (2004) placed all three species in the subgenus Hippotigris. A 2013 phylogenetic study found that the plains zebra is more closely related to Grévy's zebras than mountain zebras. The extinct quagga was originally classified as a distinct species. Later genetic studies have placed it as the same species as the plains zebra, either a subspecies or just the southernmost population. Molecular evidence supports zebras as a monophyletic lineage.
Equus originated in North America and direct paleogenomic sequencing of a 700,000-year-old middle Pleistocene horse metapodial bone from Canada implies a date of 4.07 million years ago (mya) for the most recent common ancestor of the equines within a range of 4.0 to 4.5 mya. Horses split from asses and zebras around this time and equines colonised Eurasia and Africa around 2.1–3.4 mya. Zebras and asses diverged from each other close to 2 mya. The mountain zebra diverged from the other species around 1.6 mya and the plains and Grévy's zebra split 1.4 mya.
A 2017 mitochondrial DNA study placed the Eurasian Equus ovodovi and the subgenus Sussemionus lineage as closer to zebras than to asses. However, other studies disputed this placement, finding the Sussemionus lineage basal to the zebra+asses group, but suggested that the Sussemionus lineage may have received gene flow from zebras.
The cladogram of Equus below is based on Vilstrup and colleagues (2013) and Jónsson and colleagues (2014):
Extant species Name Dimensions Description Distribution Subspecies Chromosomes Image Grévy's zebra (Equus grevyi)Body length: Tail length: Shoulder height: Weight: .Thin, elongated skull, robust neck and conical ears; narrow striping pattern with concentric rump stripes, white belly and tail base and white line around the ashy muzzle. Eastern Africa including the Horn; arid and semiarid grasslands and shrublands.Monotypic46Plains zebra (Equus quagga)Body length: Tail length: Shoulder height: Weight: .Thick bodied with relatively short legs and an obtusely shaped skull profile with a protruding forehead and a more recessed nose area; broad stripes, horizontal on the rump, with northern populations having more extensive striping while populations further south have whiter legs and bellies and more brown "shadow" stripes while the snout is black.Eastern and southern Africa; savannahs, grasslands and open woodlands.6 or monotypic44Mountain zebra (Equus zebra)Body length: Tail length: Shoulder height: Weight: .Eye sockets more circular and positioned farther back, a squarer nuchal crest, dewlap present under neck and compact hooves; stripes intermediate in width between the other species, with gridiron and horizontal stripes on the rump, while the belly is white and the black muzzle is lined with chestnut or orange.Southwestern Africa; mountains, rocky uplands and Karoo shrubland.232
Fossil record
In addition to the three living species, some fossil zebras and relatives have also been identified. E. oldowayensis is identified from remains in Olduvai Gorge dating to 1.8 mya. Fossil skulls of E. mauritanicus from Algeria which date to around 1 mya appears to show affinities with the plains zebra. E. capensis, known as the Cape zebra, appeared around 2 mya and lived throughout southern and eastern Africa.
Hybridisation
Fertile hybrids have been reported in the wild between plains and Grévy's zebra. Hybridisation has also been recorded between the plains and mountain zebra, though it is possible that these are infertile due to the difference in chromosome numbers between the two species. Captive zebras have been bred with horses and donkeys; these are known as zebroids. A zorse is a cross between a zebra and a horse; a zonkey, between a zebra and a donkey; and a zoni, between a zebra and a pony. Zebroids are often born sterile with dwarfism.
Characteristics
As with all wild equines, zebras have barrel-chested bodies with tufted tails, elongated faces and long necks with long, erect manes. Their thin legs are each supported by a spade-shaped toe covered in a hard hoof. Their dentition is adapted for grazing; they have large incisors that clip grass blades and rough molars and premolars well suited for grinding. Males have spade-shaped canines, which can be used as weapons in fighting. The eyes of zebras are at the sides and far up the head, which allows them to look over the tall grass while feeding. Their moderately long, erect ears are movable and can locate the source of a sound.
Unlike horses, zebras and asses have chestnut callosities present only on their front legs. In contrast to other living equines, zebras have longer front legs than back legs. Diagnostic traits of the zebra skull include: its relatively small size with a straight dorsal outline, protruding eye sockets, narrower rostrum, less conspicuous postorbital bar, separation of the metaconid and metastylid of the tooth by a V-shaped canal and rounded enamel wall.
Stripes
Zebras are easily recognised by their bold black-and-white striping patterns. The coat appears to be white with black stripes, as indicated by the belly and legs when unstriped, but the skin is black. Young or foals are born with brown and white coats, and the brown darkens with age. A dorsal stripe acts as the backbone for vertical stripes along the sides, from the head to the rump. On the snout they curve toward the nostrils, while the stripes above the front legs split into two branches. On the rump, they develop into species-specific patterns. The stripes on the legs, ears and tail are separate and horizontal.
Striping patterns are unique to an individual and heritable. During embryonic development, the stripes appear at eight months, but the patterns may be determined at three to five weeks. For each species there is a point in embryonic development where the stripes are perpendicular to the dorsal line and spaced apart. However, this happens at three weeks of development for the plains zebra, four weeks for the mountain zebra, and five for Grévy's zebra. The difference in timing is thought to be responsible for the differences in the striping patterns of the different species.
Various abnormalities of the patterns have been documented in plains zebras. In "melanistic" zebras, dark stripes are highly concentrated on the torso but the legs are whiter. "Spotted" individuals have broken up black stripes around the dorsal area. There have even been morphs with white spots on dark backgrounds. Striping abnormalities have been linked to inbreeding. Albino zebras have been recorded in the forests of Mount Kenya, with the dark stripes being blonde. The quagga had brown and white stripes on the head and neck, brown upper parts and a white belly, tail and legs.
Function The function of stripes in zebras has been discussed among biologists since at least the 19th century. Popular hypotheses include the following: The crypsis hypothesis suggests that the stripes allow the animal to blend in with its environment or break up its outline. This was the earliest hypothesis and proponents argued that the stripes were particularly suited for camouflage in tall grassland and woodland habitat. Alfred Wallace also wrote in 1896 that stripes make zebras less noticeable at night. Biologist Tim Caro notes that zebras graze in open habitat and do not behave cryptically, being noisy, fast, and social and do not freeze when a predator is near. In addition, the camouflaging stripes of woodland living ungulates like bongos and bushbucks are much less vivid with less contrast with the background colour. A 1987 Fourier analysis study concluded that the patterns of zebra stripes do not line up with their environment, while a 2014 study of wild equine species and subspecies could not find any correlations between striping patterns and woodland habitats. Melin and colleagues (2016) found that lions and hyenas do not appear to perceive the stripes when they are a certain distance away at daytime or nighttime, thus making the stripes useless in blending in except when the predators are close enough by which they could smell or hear their target. They also found that the stripes do not make the zebra less noticeable than solidly coloured herbivores on the open plains. They suggested that stripes may give zebras an advantage in woodlands, as the dark stripes could line up with the outlines of tree branches and other vegetation.