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Even-toed ungulates
Temporal range: Lower Eocene
Gabelbock fws 1b.jpg
Pronghorn (Antilocapra americana)
Scientific classification

Owen, 1848

The even-toed ungulates are mammals of the order Artiodactyla.

Even-toed ungulates have an even number of toes: two to four. For example, camelids or animals of the Giraffidae family have two toes, but hippopotami have four toes.

Excluding whales, the roughly 220 artiodactyl species include pigs, peccaries, hippopotamuses, cetaceans, camels, llamas, alpacas, mouse deer, deer, giraffes, antelopes, sheep, goats, and cattle, many of which are of great dietary, economic, and cultural importance to humans.


The Artiodactyla
Even-toed ungulates. While cetaceans such as whales and dolphins are not normal artiodactyls, they belong to the clade; the Artiodactyla are sometimes termed the Cetartiodactyla to reflect this


Tragulus javanicus
The mouse deer is the smallest even-toed ungulate.

Artiodactyls are generally quadrupeds. Two major body types are known: Suinas and hippopotamuses are characterized by a stocky body, short legs, and a large head; camels and ruminants, though, have a more slender build and lanky legs. Size varies considerably; the smallest member, the mouse deer, often reaches a body length of only 45 cm (18 in) and a weight of 1.5 kilograms (3.3 lb). The largest member, the hippopotamus, can grow up to 5 meters (16 ft) in length and weigh 4.5 metric tons (5.0 short tons), and the giraffe can grow to be 5.5 meters (18 ft) tall and 4.7 meters (15 ft) in body length. All even-toed ungulates display some form of sexual dimorphism: the males are consistently larger and heavier than the females. In deer, only the males boast antlers, and the horns of bovines are usually small or not present in females. Male Indian antelopes have a much darker coat than females.

Almost all even-toed ungulates have fur, with an exception being the nearly hairless hippopotamus. Fur varies in length and coloration depending on the habitat. Species in cooler regions can shed their coat. Camouflaged coats come in colors of yellow, gray, brown, or black tones.


Even-toed ungulates bear their name because they have an even number of toes (two or four) – in some peccaries, the hind legs have a reduction in the number of toes to three. The central axis of the leg is between the third and fourth toe. The first toe is missing in modern artiodactyls, and can only be found in now-extinct genera. The second and fifth toes are designed differently between species: in the hippos, they are directed forward and fully functional; for the other even-toed ungulates, they face backwards or are completely reduced. For pigs and deer, the toes are still in contact with soft, muddy ground and increase the contact surface area. In most cases, however, they no longer touch the ground. In some groups, like the camels and giraffes, regression has progressed so far that the second and fifth toe are not even present.

When camels have only two toes present, the claws are transformed into nails (while both are made of keratin, claws are curved and pointed while nails are flat and dull). These claws consist of three parts: the plate (top and sides), the sole (bottom), and the bale (rear). In general, the claws of the forelegs are wider and blunter than those of the hind legs, and the gape is farther apart. Aside from camels, all even-toed ungulates put just the tip of the foremost phalanx on the ground.

Hand skeletons with Artiodactyls highlighted
Hand skeletons of various mammals, left to right: orangutan, dog, pig, cow, tapir, and horse. Those of the artiodactyls (pig and cow) are highlighted.

In even-toed ungulates, the bones of the stylopodium (upper arm or thigh bone) and zygopodiums (tibia and fibula) are usually elongated. The muscles of the limbs are predominantly localized, which ensures that artiodactyls often have very slender legs. A clavicle is never present, and the scapula is very agile and swings back and forth for added mobility when running. The special construction of the legs causes the legs to be unable to rotate, which allows for greater stability when running at high speeds. In addition, many smaller artiodactyls have a very flexible body, contributing to their speed by increasing their stride length.


Many even-toed ungulates have a relatively large head. The skull is elongated and rather narrow; the frontal bone is enlarged near the back and displaces the parietal bone, which forms only part of the side of the cranium (especially in ruminants).

Horns and antlers

Gemsbok Kgalagadi
Outgrowths of the frontal bone characterize most forehead weapons carriers, such as the gemsbok and its horns.

Four families of even-toed ungulates have cranial appendages. These Pecora, (with the exception of the musk deer), have one of four types of cranial appendages: true horns, antlers, ossicones, or pronghorns.

True horns have a bone core that is covered in a permanent sheath of keratin, and are found only in the bovids. Antlers are bony structures that are shed and replaced each year; they are found in deer (members of the family Cervidae). They grow from a permanent outgrowth of the frontal bone called the pedicle and can be branched, as in the white-tailed deer (Odocoileus virginianus), or palmate, as in the moose (Alces alces). Ossicones are permanent bone structures that fuse to the frontal or parietal bones during an animal's life and are found only in the Giraffidae. Pronghorns, while similar to horns in that they have keratinous sheaths covering permanent bone cores, are deciduous.

All these cranial appendages can serve for posturing, battling for mating privilege, and for defense. In almost all cases, they are sexually dimorphic, and often found only on the males.


The canines of Suinas develop into tusks.

There are two trends in terms of teeth within Artiodactyla. The Suina and hippopotamuses have a relatively large number of teeth (with some pigs having 44); their dentition is more adapted to a squeezing mastication, which is characteristic of omnivores. Camels and ruminants have fewer teeth; there is often a yawning diastema, a designated gap in the teeth where the molars are aligned for crushing plant matter.

The incisors are often reduced in ruminants, and are completely absent in the upper jaw. The canines are enlarged and tusk-like in the Suina, and are used for digging in the ground and for defense. In ruminants, the males' upper canines are enlarged and used as a weapon in certain species (mouse deer, musk deer, water deer); species with frontal weapons are usually missing the upper canines. The lower canines of ruminants resemble the incisors, so that these animals have eight uniform teeth in the frontal part of the lower jaw.

The molars of porcine have only a few bumps. In contrast, the camels and ruminants have bumps that are crescent-shaped cusps (selenodont).


Artiodactyls have a well-developed sense of smell and sense of hearing. Unlike many other mammals, they have a poor sense of sight – moving objects are much easier to see than stationary ones. Similar to many other prey animals, their eyes are on the sides of the head, giving them an almost panoramic view.


Distribution and habitat

Artiodactyls are native to almost all parts of the world, with the exception of Oceania and Antarctica. Humans have introduced different artiodactyls worldwide as hunting animals. Artiodactyls inhabit almost every habitat, from tropical rain forests and steppes to deserts and high mountain regions. The greatest biodiversity prevails in open habitats such as grasslands and open forests.

Social behavior

Impalas and Giraffes Benh
Artiodactyls, like impalas and giraffes, live in groups.

The social behavior of even-toed ungulates varies from species to species. Generally, there is a tendency to merge into larger groups, but some live alone or in pairs. Species living in groups often have a hierarchy, both among males and females. Some species also live in harem groups, with one male, several females, and their common offspring. In other species, the females and juveniles stay together, while males are solitary or live in bachelor groups and seek out females only during mating season.

Many artiodactyls are territorial and mark their territory, for example, with glandular secretions or urine. In addition to year-round sedentary species, there are animals that migrate seasonally.

There are diurnal, crepuscular, and nocturnal artiodactyls. Some species' pattern of wakefulness varies with season or habitat.

Reproduction and life expectancy

Wildebeest calves, Kruger
Most artiodactyls, such as the wildebeest, are born with hair.

Generally, even-toed ungulates tend to have long gestation periods, smaller litter sizes, and more highly developed newborns. As with many other mammals, species in temperate or polar regions have a fixed mating season, while those in tropical areas breed year-round.

The length of the gestation period varies from four to five months for porcine, deer, and musk deer; six to ten months for hippos, deer, and bovines; ten to thirteen months with camels; and fourteen to fifteen months with giraffes. Most deliver one or two babies, but some pigs can deliver up to ten.

The newborns are precocial (born relatively mature) and come with open eyes and are hairy (with the exception of the hairless hippos). Juvenile deer and pigs have striped or spotted coats; the pattern disappears as they grow older. The juveniles of some species spend their first weeks with their mother in a safe location, where others may be running and following the herd within a few hours or days.

The life expectancy is typically twenty to thirty years; as in many mammals, smaller species often have a shorter lifespan than larger species. The artiodactyls with the longest lifespans are the hippos, cows, and camels, which can live 40 to 50 years.


Artiodactyls have different natural predators depending on their size and habitat. There are several carnivores that would prey on such animals, including as large cats (e.g., lions) and bears. Other predators are crocodiles, wolves, large raptors, and for small species and young animals, large snakes.

Interactions with humans


Ardiak bazkan ari dira
Some artiodactyls, like sheep, have been domesticated for thousands of years

Artiodactyls have been hunted by primitive humans for various reasons: for meat or fur, as well as to use their forehead weapons, bones, and teeth as weapons or tools. Their domestication began around the 8000 BCE. To date, humans have domesticated goats, sheep, cattle, camels, llamas, alpacas, and pigs. Initially, livestock was used primarily for food, but they began being used for work activities around 3000 BCE. Clear evidence exists of antelope being used for food 2 million years ago in the Olduvai Gorge, part of the Great Rift Valley. Cro-Magnons relied heavily on reindeer for food, skins, tools, and weapons; with dropping temperatures and increased reindeer numbers at the end of the Pleistocene, they became the prey of choice. Reindeer remains accounted for 94% of bones and teeth found in a cave above the Céou River that was inhabited around 12,500 years ago.

Today, artiodactyls are kept primarily for their meat, milk, and wool, fur, or hide for clothing. Domestic cattle, the water buffalo, the yak, and camels are used for work, as rides, or as pack animals

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