Even-toed ungulate

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Even-toed ungulates
Temporal range: 55.4–0  Ma
Early EoceneHolocene
The Cetartiodactyla.jpg
Clockwise from center: American bison (Bison bison), dromedary (Camelus dromedarius), wild boar (Sus scrofa), orca (Orcinus orca), red deer (Cervus elaphus), and giraffe (Genus: Giraffa)
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Magnorder: Boreoeutheria
Superorder: Laurasiatheria
Clade: Scrotifera
Grandorder: Ferungulata
Clade?: Ungulata
Order: Artiodactyla
Owen, 1848


The even-toed ungulates (Artiodactyla /ˌɑːrtiˈdæktɪlə/ , from Ancient Greek ἄρτιος, ártios 'even',andδάκτυλος, dáktylos 'finger, toe') are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing posteriorly. By contrast, odd-toed ungulates bear weight on an odd number of the five toes. Another difference between the two is that many other even-toed ungulates (except for Suina) digest plant cellulose in one or more stomach chambers rather than in their intestine as the odd-toed ungulates do.


The advent of molecular biology, along with new fossil discoveries, found that Cetaceans (whales, dolphins, and porpoises) fall within this taxonomic branch, being most closely related to hippopotamuses. Some modern taxonomists thus apply the name Cetartiodactyla /sɪˌtɑːrtiˈdæktɪlə/ to this group, while others opt to include cetaceans within the existing name of Artiodactyla.

The roughly 270 land-based even-toed ungulate species include pigs, peccaries, hippopotamuses, antelopes, deer, giraffes, camels, llamas, alpacas, sheep, goats, and cattle. Many are herbivores, but suids are omnivorous, while cetaceans are almost exclusively carnivorous. Many of these are of great dietary, economic, and cultural importance to humans.

Evolutionary history

The oldest fossils of even-toed ungulates date back to the early Eocene (about 53 million years ago). Since these findings almost simultaneously appeared in Europe, Asia, and North America, it is very difficult to accurately determine the origin of artiodactyls. The fossils are classified as belonging to the family Dichobunidae; their best-known and best-preserved member is Diacodexis . [1] These were small animals, some as small as a hare, with a slim build, lanky legs, and a long tail. Their hind legs were much longer than their front legs. The early to middle Eocene saw the emergence of the ancestors of most of today's mammals. [2]

Entelodonts were stocky animals with a large head, and were characterized by bony bumps on the lower jaw. Entelodon Sp Illustration.jpg
Entelodonts were stocky animals with a large head, and were characterized by bony bumps on the lower jaw.

Two formerly widespread, but now extinct, families of even-toed ungulates were Entelodontidae and Anthracotheriidae. Entelodonts existed from the middle Eocene to the early Miocene in Eurasia and North America. They had a stocky body with short legs and a massive head, which was characterized by two humps on the lower jaw bone. Anthracotheres had a large, porcine (pig-like) build, with short legs and an elongated muzzle. This group appeared in the middle Eocene up until the Pliocene, and spread throughout Eurasia, Africa, and North America. Anthracotheres are thought to be the ancestors of hippos, and, likewise, probably led a similar aquatic lifestyle. Hippopotamuses appeared in the late Miocene and occupied Africa and Asia—they never got to the Americas. [2]

The camels (Tylopoda) were, during large parts of the Cenozoic, limited to North America; early forms like Cainotheriidae occupied Europe. Among the North American camels were groups like the stocky, short-legged Merycoidodontidae. They first appeared in the late Eocene and developed a great diversity of species in North America. Only in the late Miocene or early Pliocene did they migrate from North America into Eurasia. The North American varieties became extinct around 10,000 years ago.

Suina (including pigs) have been around since the Eocene. In the late Eocene or the Oligocene, two families stayed in Eurasia and Africa; the peccaries, which became extinct in the Old World, exist today only in the Americas.

Sivatherium was a relative of giraffes with deer-like forehead weapons. MEPAN Sivatherium.jpg
Sivatherium was a relative of giraffes with deer-like forehead weapons.

South America was settled by even-toed ungulates only in the Pliocene, after the land bridge at the Isthmus of Panama formed some three million years ago. With only the peccaries, lamoids (or llamas), and various species of capreoline deer, South America has comparatively fewer artiodactyl families than other continents, except Australia, which has no native species.

Taxonomy and phylogeny

Richard Owen coined the term "even-toed ungulate". Thomas Herbert Maguire - Richard Owen 1850.png
Richard Owen coined the term "even-toed ungulate".

The classification of artiodactyls was hotly debated because the ocean-dwelling cetaceans evolved from the land-dwelling even-toed ungulates. Some semiaquatic even-toed ungulates (hippopotamuses) are more closely related to the ocean-dwelling cetaceans than to the other even-toed ungulates.

This makes the Artiodactyla as traditionally defined a paraphyletic taxon, since it includes animals descended from a common ancestor, but does not include all of its descendants. Phylogenetic classification only recognizes monophyletic taxa; that is, groups that descend from a common ancestor and include all of its descendants. To address this problem, the traditional order Artiodactyla and infraorder Cetacea are sometimes subsumed into the more inclusive Cetartiodactyla taxon. [3] An alternative approach is to include both land-dwelling even-toed ungulates and ocean-dwelling cetaceans in a revised Artiodactyla taxon. [2]


Research history

Molecular and morphological studies confirmed that cetaceans are the closest living relatives of hippopotamuses. Humpback Whale underwater shot.jpg
Molecular and morphological studies confirmed that cetaceans are the closest living relatives of hippopotamuses.

In the 1990s, biological systematics used not only morphology and fossils to classify organisms, but also molecular biology. Molecular biology involves sequencing an organism's DNA and RNA and comparing the sequence with that of other living beings—the more similar they are, the more closely they are related. Comparison of even-toed ungulate and cetaceans genetic material has shown that the closest living relatives of whales and hippopotamuses is the paraphyletic group Artiodactyla.

Dan Graur and Desmond Higgins were among the first to come to this conclusion, and included a paper published in 1994. [5] However, they did not recognize hippopotamuses and classified the ruminants as the sister group of cetaceans. Subsequent studies established the close relationship between hippopotamuses and cetaceans; these studies were based on casein genes, [6] SINEs, [7] fibrinogen sequences, [8] cytochrome and rRNA sequences, [3] [9] IRBP (and vWF) gene sequences, [10] adrenergic receptors, [11] and apolipoproteins. [12]

In 2001, the fossil limbs of a Pakicetus (amphibioid cetacean the size of a wolf) and Ichthyolestes (an early whale the size of a fox) were found in Pakistan. They were both archaeocetes ("ancient whales") from about 48 million years ago (in the Eocene). These findings showed that archaeocetes were more terrestrial than previously thought, and that the special construction of the talus (ankle bone) with a double-rolled joint surface,[ clarification needed ] previously thought to be unique to even-toed ungulates, were also in early cetaceans. [13] The mesonychids, another type of ungulate, did not show this special construction of the talus, and thus was concluded to not have the same ancestors as cetaceans.

Hippos are a geologically young group, which raises questions about their origin. Hippopotamus amphibius.JPG
Hippos are a geologically young group, which raises questions about their origin.

The oldest cetaceans date back to the early Eocene (53 million years ago), whereas the oldest known hippopotamus dates back only to the Miocene (15 million years ago). The hippopotamids are descended from the anthracotheres, a family of semiaquatic and terrestrial artiodactyls that appeared in the late Eocene, and are thought to have resembled small- or narrow-headed hippos. Research is therefore focused on anthracotheres (family Anthracotheriidae); one dating from the Eocene to Miocene was declared to be "hippo-like" upon discovery in the 19th century. A study from 2005 showed that the anthracotheres and hippopotamuses have very similar skulls, but differed in the adaptations of their teeth. It was nevertheless believed that cetaceans and anthracothereres descended from a common ancestor, and that hippopotamuses developed from anthracotheres. A study published in 2015 was able to confirm this, but also revealed that hippopotamuses were derived from older anthracotherians. [9] [14] The newly introduced genus Epirigenys from eastern Africa is thus the sister group of hippos.

Morphological classification of Artiodactyla

Linnaeus postulated a close relationship between camels and ruminants as early as the mid-1700s.[ citation needed ] Henri de Blainville recognized the similar anatomy of the limbs of pigs and hippos,[ when? ] and British zoologist Richard Owen coined the term "even-toed ungulates" and the scientific name "Artiodactyla" in 1848.[ citation needed ]

Internal morphology (mainly the stomach and the molars) were used for classification. Suines (including pigs) and hippopotamuses have molars with well-developed roots and a simple stomach that digests food. Thus, they were grouped together as non-ruminants (Porcine). All other even-toed ungulates have molars with a selenodont construction (crescent-shaped cusps) and have the ability to ruminate, which requires regurgitating food and re-chewing it. Differences in stomach construction indicated that rumination evolved independently between tylopods and ruminants; therefore, tylopods were excluded from Ruminantia.

The taxonomy that was widely accepted by the end of the 20th century was: [15] [ full citation needed ]

Even-toed ungulates

  Suidae Recherches pour servir a l'histoire naturelle des mammiferes (Pl. 80) (white background).jpg

  Hippopotamidae Voyage en Abyssinie Plate 2 (white background).jpg


  Tylopoda Cladogram of Cetacea within Artiodactyla (Camelus bactrianus).png


  Tragulidae Tragulus napu - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg

  Pecora Walia ibex illustration white background.png

Morphological classification of Cetacea

The mesonychids were long considered ancestors of whales. Mesonyx.jpg
The mesonychids were long considered ancestors of whales.

Modern cetaceans are highly adapted sea creatures which, morphologically, have little in common with land mammals; they are similar to other marine mammals, such as seals and sea cows, due to convergent evolution. However, they evolved from originally terrestrial mammals. The most likely ancestors were long thought to be mesonychids—large, carnivorous animals from the early Cenozoic (Paleocene and Eocene), which had hooves instead of claws on their feet. Their molars were adapted to a carnivorous diet, resembling the teeth in modern toothed whales, and, unlike other mammals, have a uniform construction.[ citation needed ]

The suspected relations can be shown as follows: [14] [16] [ page needed ]


  Artiodactyla Walia ibex illustration white background.png


  Mesonychia Synoplotherium112DB.jpg

  Cetacea Bowhead-Whale1 (16273933365).jpg

Inner systematics

Molecular findings and morphological indications suggest that artiodactyls as traditionally defined are paraphyletic with respect to cetaceans. Cetaceans are deeply nested within the former; the two groups together form a monophyletic taxon, for which the name Cetartiodactyla is sometimes used. Modern nomenclature divides Artiodactyla (or Cetartiodactyla) in four subordinate taxa: camelids (Tylopoda), pigs and peccaries (Suina), ruminants (Ruminantia), and hippos plus whales (Whippomorpha).

The presumed lineages within Artiodactyla can be represented in the following cladogram: [17] [18] [19] [20] [21]


  Tylopoda (camels) Cladogram of Cetacea within Artiodactyla (Camelus bactrianus).png


  Suina (pigs) Recherches pour servir a l'histoire naturelle des mammiferes (Pl. 80) (white background).jpg

   Ruminantia  (ruminants)  

  Tragulidae (mouse deer) Tragulus napu - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg

  Pecora (horn bearers) Walia ibex illustration white background.png


  Hippopotamidae (hippopotamuses) Voyage en Abyssinie Plate 2 (white background).jpg

  Cetacea (whales) Bowhead-Whale1 (16273933365).jpg

  (or  Whippomorpha)  
Camels are now considered a sister group of Artiofabula. Camelus dromedarius in Singapore Zoo.JPG
Camels are now considered a sister group of Artiofabula.
The pronghorn is the only extant antilocaprid. Antilocapra americana.jpg
The pronghorn is the only extant antilocaprid.

The four summarized Artiodactyla taxa are divided into ten extant families: [22]

Although deer, musk deer, and pronghorns have traditionally been summarized as cervids (Cervioidea), molecular studies provide different—and inconsistent—results, so the question of phylogenetic systematics of infraorder Pecora (the horned ruminants) for the time being, cannot be answered.

Reconstruction of Indohyus Indohyus BW.jpg
Reconstruction of Indohyus


Blue duiker (Philantomba monticola) skeleton on display at the Museum of Osteology. Blue Duiker skeleton.jpg
Blue duiker ( Philantomba monticola) skeleton on display at the Museum of Osteology.

Artiodactyls are generally quadrupeds. Two major body types are known: Suinids 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 centimeters (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 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.


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

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 adapted differently between species:

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). [24] 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 they are farther apart. Aside from camels, all even-toed ungulates put just the tip of the foremost phalanx on the ground. [25]

Diagrams of hand skeletons of various mammals, left to right: orangutan, dog, pig, cow, tapir, and horse. Highlighted are the even-toed ungulates pig and cow. Hand skeletons with Artiodactyls highlighted.png
Diagrams of hand skeletons of various mammals, left to right: orangutan, dog, pig, cow, tapir, and horse. Highlighted are the even-toed ungulates pig and cow.

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

Outgrowths of the frontal bone characterize most forehead weapons carriers, such as the gemsbok and its horns. Gemsbok Kgalagadi.jpg
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. [26]

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.[ clarification needed ] [27]

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. One exception is the species Rangifer tarandus, known as reindeer in Europe or caribou in North America, where both sexes can grow antlers yearly, though the females' antlers are typically smaller and not always present.


The canines of Suinas develop into tusks. Hirscheber1a.jpg
The canines of Suinas develop into tusks.
Dental formula I C P M

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.

Digestive system

Pigs (such as this warthog) have a simple sack-shaped stomach. Warthog Face 001.jpg
Pigs (such as this warthog) have a simple sack-shaped stomach.
As with all ruminants, deer have such a multi-chambered stomach, which is used for better digesting plant food. Rothirsch.jpg
As with all ruminants, deer have such a multi-chambered stomach, which is used for better digesting plant food.

The ruminants (Ruminantia) ruminate their food—they regurgitate and re-chew it. Ruminants' mouths often have additional salivary glands, and the oral mucosa is often heavily calloused to avoid injury from hard plant parts and to allow easier transport of roughly chewed food. Their stomachs are divided into three to four sections: the rumen, the reticulum, the omasum, and the abomasum. [28] After the food is ingested, it is mixed with saliva in the rumen and reticulum and separates into layers of solid versus liquid material. The solids lump together to form a bolus (also known as the cud); this is regurgitated by reticular contractions while the glottis is closed. When the bolus enters the mouth, the fluid is squeezed out with the tongue and re-swallowed. The bolus is chewed slowly to completely mix it with saliva and to break it down. Ingested food passes to the "fermentation chamber" (rumen and reticulum), where it is kept in continual motion by rhythmic contractions. Cellulytic microbes (bacteria, protozoa, and fungi) produce cellulase, which is needed to break down the cellulose found in plant material. [28] This form of digestion has two advantages: plants that are indigestible to other species can be digested and used, and the duration of the actual food consumption shortened; the animal spends only a short time out in the open with his head to the ground—rumination can take place later, in a sheltered area. [29]

Tylopoda (camels, llamas, and alpacas) and chevrotains have three-chambered stomachs, while the rest of Ruminantia have four-chambered stomachs. The handicap of a heavy digestive system has increased selective pressure towards limbs that allow the animal to quickly escape predators. [30] Most species within Suina have a simple two-chambered stomach that allows an omnivorous diet. The babirusa, however, is an herbivore, [28] and has extra maxillary teeth to allow for proper mastication of plant material. Most of the fermentation occurs with the help of cellulolytic microorganisms within the caecum of the large intestine. Peccaries have a complex stomach that contains four compartments. [29] Their fore stomach has fermentation carried out by microbes and has high levels of volatile fatty acid; it has been proposed that their complex fore stomach is a means to slow digestive passage and increase digestive efficiency. [29] Hippopotamuses have three-chambered stomachs and do not ruminate. They consume around 68 kilograms (150 lb) of grass and other plant matter each night. They may cover distances up to 32 kilometers (20 mi) to obtain food, which they digest with the help of microbes that produce cellulase. Their closest living relatives, the whales, are obligate carnivores.

Unlike other even-toed ungulates, pigs have a simple sack-shaped stomach. [28] Some artiodactyla, such as white-tailed deer, lack a gall bladder. [31]

The Japanese serow has glands in the eyes that are clearly visible Lightmatter japanese serows.jpg
The Japanese serow has glands in the eyes that are clearly visible

Genitourinary system

The penises of even-toed ungulates have an S-shape at rest and lie in a pocket under the skin on the belly. The corpora cavernosa are only slightly developed; and an erection mainly causes this curvature to extend, which leads to an extension, but not a thickening, of the penis. Cetaceans have similar penises. [32] In some even-toed ungulates, the penis contains a structure called the urethral process. [33] [34] [35]

The testicles are located in the scrotum and thus outside the abdominal cavity. The ovaries of many females descend—as testicles descend of many male mammals—and are close to the pelvic inlet at the level of the fourth lumbar vertebra. The uterus has two horns (uterus bicornis). [32]


The number of mammary glands is variable and correlates, as in all mammals, with litter size. Pigs, which have the largest litter size of all even-toed ungulates, have two rows of teats lined from the armpit to the groin area. In most cases, however, even-toed ungulates have only one or two pairs of teats. In some species these form an udder in the groin region.

Secretory glands in the skin are present in virtually all species and can be located in different places, such as in the eyes, behind the horns, the neck, or back, on the feet, or in the anal region.


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. [36] Artiodactyls inhabit almost every habitat, from tropical rainforests and steppes to deserts and high mountain regions. The greatest biodiversity prevails in open habitats such as grasslands and open forests.

Social behavior

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

Most artiodactyls, such as the wildebeest, are born with hair. Wildebeest calves, Kruger.jpg
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. They carry out polygynous mating behavior, meaning a male mates with several females and suppresses all competition.

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.

Predators and parasites

Artiodactyls have different natural predators depending on their size and habitat. There are several carnivores that prey on them, including large cats (e.g., lions) and bears. Other predators are crocodiles, wolves and dogs, large raptors, and for small species and young animals, large snakes. For cetaceans, possible predators include sharks, polar bears, and other cetaceans; in the latter is the orca, the top predator of the oceans. [37]

Parasites include nematodes, botflies, fleas, lice, or flukes, but they have debilitating effects only when the infestation is severe.[ citation needed ]

Interactions with humans


Some artiodactyls, like sheep, have been domesticated for thousands of years. Ardiak bazkan ari dira.JPG
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 bones and teeth as weapons or tools. Their domestication began around 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. [30] Clear evidence exists of antelope being used for food 2 million years ago in the Olduvai Gorge, part of the Great Rift Valley. [30] [38] 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 river Céou that was inhabited around 12,500 years ago. [39]

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. [40] [ page needed ]


The aurochs has been extinct since the 17th century. Ur-painting.jpg
The aurochs has been extinct since the 17th century.

The endangerment level of each even-toed ungulate is different. Some species are synanthropic (such as the wild boar) and have spread into areas that they are not indigenous to, either having been brought as farm animals or having run away as people's pets. Some artiodactyls also benefit from the fact that their predators (e.g. the Tasmanian tiger) were severely decimated by ranchers, who saw them as competition. [36]

Conversely, many artiodactyls have declined significantly in numbers, and some have even gone extinct, largely due to over-hunting, and, more recently, habitat destruction. Extinct species include several gazelles, the aurochs, the Malagasy hippopotamus, the bluebuck, and Schomburgk's deer. Two species, the Scimitar-horned oryx and Père David's deer, are extinct in the wild. Fourteen species are considered critically endangered, including the addax, the kouprey, the Bactrian camel, Przewalski's gazelle, the saiga, and the pygmy hog. Twenty-four species are considered endangered. [41] [42]

See also

Related Research Articles

<span class="mw-page-title-main">Ungulate</span> Group of animals that use the tips of their toes or hooves to walk on

Ungulates are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. Living ungulates are divided into two orders: the odd-toed ungulates (Perissodactyla) including horses, rhinoceroses, and tapirs; and even-toed ungulates (Artiodactyla) such as cattle, pigs, giraffes, camels, sheep, deer, and hippopotamuses. Cetaceans such as whales, dolphins, and porpoises are also classified as even-toed ungulates, although they do not have hooves. Most terrestrial ungulates use the hoofed tips of their toes to support their body weight while standing or moving. Two other orders of ungulates, Notoungulata and Litopterna, both native to South America, became extinct at the end of the Pleistocene, around 12,000 years ago.

<span class="mw-page-title-main">Hippopotamidae</span> Family of mammals

Hippopotamidae is a family of stout, naked-skinned, and semiaquatic artiodactyl mammals, possessing three-chambered stomachs and walking on four toes on each foot. While they resemble pigs physiologically, their closest living relatives are the cetaceans. They are formally referred to as hippopotamids.

<span class="mw-page-title-main">Pygmy hippopotamus</span> Small species of hippopotamus from West Africa

The pygmy hippopotamus or pygmy hippo is a small hippopotamid which is native to the forests and swamps of West Africa, primarily in Liberia, with small populations in Sierra Leone, Guinea, and Ivory Coast. It has been extirpated from Nigeria.

<span class="mw-page-title-main">Tylopoda</span> Suborder of mammals

Tylopoda is a suborder of terrestrial herbivorous even-toed ungulates belonging to the order Artiodactyla. They are found in the wild in their native ranges of South America and Asia, while Australian feral camels are introduced. The group has a long fossil history in North America and Eurasia. Tylopoda appeared during the Eocene around 50 million years ago.

<span class="mw-page-title-main">Evolution of cetaceans</span>

The evolution of cetaceans is thought to have begun in the Indian subcontinent from even-toed ungulates 50 million years ago (mya) and to have proceeded over a period of at least 15 million years. Cetaceans are fully aquatic marine mammals belonging to the order Artiodactyla and branched off from other artiodactyls around 50 mya. Cetaceans are thought to have evolved during the Eocene, the second epoch of the present-extending Cenozoic Era. Molecular and morphological analyses suggest Cetacea share a relatively recent closest common ancestor with hippopotami and that they are sister groups. Being mammals, they surface to breathe air; they have 5 finger bones (even-toed) in their fins; they nurse their young; and, despite their fully aquatic life style, they retain many skeletal features from their terrestrial ancestors. Research conducted in the late 1970s in Pakistan revealed several stages in the transition of cetaceans from land to sea.

<span class="mw-page-title-main">Entelodont</span> An extinct family of pig-like omnivores from North America and Eurasia

Entelodontidae is a extinct family of pig-like artiodactyls which inhabited the Northern Hemisphere from the late Eocene to the Middle Miocene epochs, about 38-19 million years ago. Their large heads, low snouts, narrow gait, and proposed omnivorous diet inspires comparisons to suids and tayassuids (peccaries), and historically they have been considered closely related to these families purely on a morphological basis. However, studies which combine morphological and molecular (genetic) data on artiodactyls instead suggest that entelodonts are cetancodontamorphs, more closely related to hippos and cetaceans through their resemblance to Pakicetus, than to basal pigs like Kubanochoerus and other ungulates.

<span class="mw-page-title-main">Mesonychid</span> Extinct taxon of carnivorous ungulates

Mesonychia is an extinct taxon of small- to large-sized carnivorous ungulates related to artiodactyls. Mesonychids first appeared in the early Paleocene, went into a sharp decline at the end of the Eocene, and died out entirely when the last genus, Mongolestes, became extinct in the early Oligocene. In Asia, the record of their history suggests they grew gradually larger and more predatory over time, then shifted to scavenging and bone-crushing lifestyles before the group became extinct.

<i>Pakicetus</i> Genus of ancient whales

Pakicetus is an extinct genus of amphibious cetacean of the family Pakicetidae, which was endemic to Pakistan during the Eocene, about 50 million years ago. It was a wolf-like animal, about 1 metre to 2 metres long, and lived in and around water where it ate fish and other small animals. The vast majority of paleontologists regard it as the most basal whale, representing a transitional stage between land mammals and whales. It belongs to the even-toed ungulates with the closest living non-cetacean relative being the hippopotamus.

<span class="mw-page-title-main">Pecora</span> Infraorder of mammals

Pecora is an infraorder of even-toed hoofed mammals with ruminant digestion. Most members of Pecora have cranial appendages projecting from their frontal bones; only two extant genera lack them, Hydropotes and Moschus. The name “Pecora” comes from the Latin word pecus, which means “horned livestock”. Although most pecorans have cranial appendages, only some of these are properly called “horns”, and many scientists agree that these appendages did not arise from a common ancestor, but instead evolved independently on at least two occasions. Likewise, while Pecora as a group is supported by both molecular and morphological studies, morphological support for interrelationships between pecoran families is disputed.

<i>Archaeotherium</i> Extinct genus of mammals

Archaeotherium is an extinct genus of entelodont artiodactyl endemic to North America during the Eocene and Oligocene epochs (35—28 mya), existing for approximately 9.1 million years. Archaeotherium fossils are most common in the White River Formation of the Great Plains, but it has also been found in the John Day Basin of Oregon and the Trans-Pecos area of Texas.

<span class="mw-page-title-main">Merycoidodontoidea</span> Extinct superfamily of mammals

Merycoidodontoidea, sometimes called "oreodonts" or "ruminating hogs", is an extinct superfamily of prehistoric cud-chewing artiodactyls with short faces and fang-like canine teeth. As their name implies, some of the better known forms were generally hog-like, and the group has traditionally been placed within the Suina, though some recent work suggests they may have been more closely related to camels. "Oreodont" means "mountain teeth", referring to the appearance of the molars. Most oreodonts were sheep-sized, though some genera grew to the size of cattle. They were heavy-bodied, with short four-toed hooves and comparatively long tails.

<span class="mw-page-title-main">Anthracotheriidae</span> Extinct family of mammals

Anthracotheriidae is a paraphyletic family of extinct, hippopotamus-like artiodactyl ungulates related to hippopotamuses and whales. The oldest genus, Elomeryx, first appeared during the middle Eocene in Asia. They thrived in Africa and Eurasia, with a few species ultimately entering North America during the Oligocene. They died out in Europe and Africa during the Miocene, possibly due to a combination of climatic changes and competition with other artiodactyls, including pigs and true hippopotamuses. The youngest genus, Merycopotamus, died out in Asia during the late Pliocene, possibly for the same reasons. The family is named after the first genus discovered, Anthracotherium, which means "coal beast", as the first fossils of it were found in Paleogene-aged coal beds in France. Fossil remains of the anthracothere genus were discovered by the Harvard University and Geological Survey of Pakistan joint research project (Y-GSP) in the well-dated middle and late Miocene deposits of the Pothohar Plateau in northern Pakistan.

<span class="mw-page-title-main">Whippomorpha</span> Suborder of mammals

Whippomorpha or Cetancodonta is a group of animals that contains all living cetaceans and hippopotamuses, as well as their extinct relatives, i.e. Entelodonts and Andrewsarchus. All Whippomorphs are descendants of the last common ancestor of Hippopotamus amphibius and Tursiops truncatus. This makes it a crown group. Whippomorpha is a suborder within the order Artiodactyla. The placement of Whippomorpha within Artiodactyla is a matter of some contention, as hippopotamuses were previously considered to be more closely related to Suidae (pigs) and Tayassuidae (peccaries). Most contemporary scientific phylogenetic and morphological research studies link hippopotamuses with cetaceans, and genetic evidence has overwhelmingly supported an evolutionary relationship between Hippopotamidae and Cetacea. Modern Whippomorphs all share a number of behavioural and physiological traits; such as a dense layer of subcutaneous fat and largely hairless bodies. They exhibit amphibious and aquatic behaviors and possess similar auditory structures.

<span class="mw-page-title-main">Cetruminantia</span> Taxonomic clade

The Cetruminantia are a clade made up of the Cetancodontamorpha and their closest living relatives, the Ruminantia.

<span class="mw-page-title-main">Protoceratidae</span> Extinct family of mammals

Protoceratidae is an extinct family of herbivorous North American artiodactyls that lived during the Eocene through Pliocene at around 46.2—4.9 Mya, existing for about 41 million years.

<i>Indohyus</i> Genus of extinct artiodactyl mammals from Eocene Epoch

Indohyus is an extinct genus of digitigrade even-toed ungulates known from Eocene fossils in Asia. This small chevrotain-like animal found in the Himalayas is one of the earliest known non-cetacean ancestors of whales.

Artiocetus is an extinct genus of early whales belonging to the family Protocetidae. It was a close relative to Rodhocetus and its tarsals indicate it resembled an artiodactyl.

<span class="mw-page-title-main">Artiofabula</span> Clade of mammals comprising pigs, cows, hippos, and whales, among others

Artiofabula is a clade made up of the Suina and the Cetruminantia. The clade was found in molecular phylogenetic analyses and contradicted traditional relationships based on morphological analyses.

<span class="mw-page-title-main">Ancodonta</span> Infraorder of mammals

Ancodonta is an infraorder of semiaquatic artiodactyl ungulates including modern hippopotamus and all mammals closer to hippos than to cetaceans (whales). Ancodonts first appeared in the Middle Eocene, with some of the earliest representatives found in fossil deposits in Southeast Asia. Throughout their evolutionary history they have occupied different browsing and grazing niches in North America, Eurasia and Africa. The last continent is notable as they were among the first laurasiatherian mammals to have migrated to Africa from Europe, where they competed with the native afrothere herbivores for the same niches. Of the nearly 50 genera that have existed, only two of them are extant – Choeropsis and Hippopotamus. The interrelationships within the ancodonts has been contended. The traditional notion is that there at minimum two families Anthracotheriidae and Hippopotamidae and were merely sister taxa. However many detailed research of the dentition among ancodonts, as well as how some anthracotheres were similar to hippos in appearance, lead the current consensus where Anthracotheriidae is paraphyletic to Hippopotamidae. Among the anthracotheres members of Bothriodontinae are among the closest to the ancestry of hippos, with the Oligocene aged Epirigenys from Lokon, Turkana, Kenya being the sister taxon to hippos. In response of this many similar clade names have been used for this clade.


  1. Theodor, Jessica M.; Erfurt, Jörg; Grégoire Métais (23 October 2007). "The earliest artiodactyls: Diacodexeidae, Dichobunidae, Homacodontidae, Leptochoeridae and Raoellidae". In Prothero, Donald R.; Foss, Scott E. (eds.). Evolution of Artiodactyls. Johns Hopkins University. pp. 32–58. ISBN   9780801887352.
  2. 1 2 3 4 Spaulding, M; O'Leary, MA; Gatesy, J (2009). Farke, Andrew Allen (ed.). "Relationships of Cetacea (Artiodactyla) Among Mammals: Increased Taxon Sampling Alters Interpretations of Key Fossils and Character Evolution". PLOS ONE. 4 (9): e7062. Bibcode:2009PLoSO...4.7062S. doi: 10.1371/journal.pone.0007062 . PMC   2740860 . PMID   19774069.
  3. 1 2 Montgelard, Claudine; Catzeflis, Francois M.; Douzery, Emmanuel (1997). "Phylogenetic relationships of artiodactyls and cetaceans as deduced from the comparison of cytochrome b and 12S rRNA mitochondrial sequences". Molecular Biology and Evolution. 14 (5): 550–559. doi: 10.1093/oxfordjournals.molbev.a025792 . PMID   9159933.
  4. Groves, Colin P.; Grubb, Peter (2011). Ungulate Taxonomy. Baltimore, Maryland: Johns Hopkins University Press. p. 25. ISBN   978-1-4214-0093-8.
  5. Graur, Dan; Higgins, Desmond G. (1994). "Molecular Evidence for the Inclusion of Cetaceans within the Order Artiodactyla" (PDF). Molecular Biology and Evolution: 357–364. Archived from the original (PDF) on 5 March 2016. Retrieved 23 August 2015.
  6. Gatesy, John; Hayashi, Cheryl; Cronin, Mathew A.; Arctander, Peter (1996). "Evidence from milk casein genes that cetaceans are close relatives of hippopotamid artiodactyls". Molecular Biology and Evolution. 13 (7): 954–963. doi: 10.1093/oxfordjournals.molbev.a025663 . PMID   8752004.
  7. Shimamura, M. (1997). "Molecular evidence from retroposons that whales form a clade within even-toed ungulates". Nature. 388 (6643): 666–670. Bibcode:1997Natur.388..666S. doi: 10.1038/41759 . PMID   9262399. S2CID   4429657. Closed Access logo transparent.svg
  8. Gatesy, John (1997). "More DNA Support for a Cetacea/Hippopotamidae Clade: The Blood-Clotting Protein Gene y-Fibrinogen". Molecular Biology and Evolution. 14 (5): 537–543. doi: 10.1093/oxfordjournals.molbev.a025790 . PMID   9159931.
  9. 1 2 Agnarsson, Ingi; May-Collado, Laura J. (2008). "The phylogeny of Cetartiodactyla: The importance of dense taxon sampling, missing data, and the remarkable promise of cytochrome b to provide reliable species-level phylogenies". Molecular Phylogenetics and Evolution. 48 (3): 964–85. doi:10.1016/j.ympev.2008.05.046. PMID   18590827.
  10. Gatesy, John; Milinkovitch, Michel; Waddell, Victor; Stanhope, Michael (1999). "Stability of Cladistic Relationships between Cetacea and Higher-Level Artiodactyl Taxa". Systematic Biology. 48 (1): 6–20. doi: 10.1080/106351599260409 . PMID   12078645.
  11. Madsen, Ole; Willemsen, Diederik; Ursing, Björn M.; Arnason, Ulfur; de Jong, Wilfried W. (2002). "Molecular Evolution of the Mammalian Alpha 2B Adrenergic Receptor". Molecular Biology and Evolution. 19 (12): 2150–2160. doi: 10.1093/oxfordjournals.molbev.a004040 . PMID   12446807.
  12. Amrine-Madsen, Heather; Koepfli, Klaus-Peter; Wayne, Robert K.; Springer, Mark S. (2003). "A new phylogenetic marker, apolipoprotein B, provides compelling evidence for eutherian relationships". Molecular Phylogenetics and Evolution. 28 (2): 225–240. doi:10.1016/s1055-7903(03)00118-0. PMID   12878460.
  13. Savage, R. J. G.; Long, M. R. (1986). Mammal Evolution: an illustrated guide . New York: Facts on File. pp.  208. ISBN   978-0-8160-1194-0.
  14. 1 2 Price, Samantha A.; Bininda-Emonds, Olaf R. P.; Gittleman, John L. (2005). "A complete phylogeny of the whales, dolphins and even-toed hoofed mammals (Cetartiodactyla)". Biological Reviews. 80 (3): 445–73. doi:10.1017/s1464793105006743. PMID   16094808. S2CID   45056197.
  15. etwa noch bei Nowak (1999) oder Hendrichs (2004)
  16. McKenna, Malcolm C.; Bell, Susan K. (1997). 'Classification of Mammals - Above the Species Level. Columbia University Press. ISBN   978-0-231-11013-6.
  17. Beck, N.R. (2006). "A higher-level MRP supertree of placental mammals". BMC Evol Biol. 6: 93. doi:10.1186/1471-2148-6-93. PMC   1654192 . PMID   17101039.
  18. O'Leary, M.A.; Bloch, J.I.; Flynn, J.J.; Gaudin, T.J.; Giallombardo, A.; Giannini, N.P.; Goldberg, S.L.; Kraatz, B.P.; Luo, Z.-X.; Meng, J.; Ni, X.; Novacek, M.J.; Perini, F.A.; Randall, Z.S.; Rougier, G.W.; Sargis, E.J.; Silcox, M.T.; Simmons, N.B.; Spaulding, M.; Velazco, P.M.; Weksler, M.; Wible, J.R.; Cirranello, A.L. (2013). "The Placental Mammal Ancestor and the Post-K-Pg Radiation of Placentals". Science. 339 (6120): 662–667. Bibcode:2013Sci...339..662O. doi:10.1126/science.1229237. hdl: 11336/7302 . PMID   23393258. S2CID   206544776.
  19. Song, S.; Liu, L.; Edwards, S.V.; Wu, S. (2012). "Resolving conflict in eutherian mammal phylogeny using phylogenomics and the multispecies coalescent model". Proceedings of the National Academy of Sciences. 109 (37): 14942–14947. Bibcode:2012PNAS..10914942S. doi: 10.1073/pnas.1211733109 . PMC   3443116 . PMID   22930817.
  20. dos Reis, M.; Inoue, J.; Hasegawa, M.; Asher, R.J.; Donoghue, P.C.J.; Yang, Z. (2012). "Phylogenomic datasets provide both precision and accuracy in estimating the timescale of placental mammal phylogeny". Proceedings of the Royal Society B: Biological Sciences. 279 (1742): 3491–3500. doi: 10.1098/rspb.2012.0683 . PMC   3396900 . PMID   22628470.
  21. Upham, N.S.; Esselstyn, J.A.; Jetz, W. (2019). "Inferring the mammal tree: Species-level sets of phylogenies for questions in ecology, evolution, and conservation". PLOS Biology. 17 (12): e3000494. doi: 10.1371/journal.pbio.3000494 . PMC   6892540 . PMID   31800571.(see e.g. Fig S10)
  22. Wilson, D. E.; Reeder, D. M., eds. (2005). Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 111–184. ISBN   978-0-8018-8221-0.
  23. Cui, P.; Ji, R.; Ding, F.; Qi, D.; Gao, H.; Meng, H.; Yu, J.; Hu, S.; Zhang, H. (2007). "A complete mitochondrial genome sequence of the wild two-humped camel (Camelus bactrianus ferus): an evolutionary history of Camelidae". BMC Genomics. 8 (1): 241. doi:10.1186/1471-2164-8-241. PMC   1939714 . PMID   17640355.
  24. "Claws Out: Things You Didn't Know About Claws". Thomson Safaris. 7 January 2014. Retrieved 24 September 2016.
  25. Salomon, Franz-Viktor; et al. (et al.) (2008). Salomon, F.-V. (ed.). musculoskeletal system. Anatomy for veterinary medicine. pp. 22–234. ISBN   978-3-8304-1075-1.
  26. DeMiguel, Daniel; Azanza, Beatriz; Morales, Jorge (2014). "Key innovations in ruminant evolution: a paleontological perspective". Integrative Zoology. 9 (4): 412–433. doi:10.1111/1749-4877.12080. PMID   24148672.
  27. Janis, C.M.; Scott, K.M. (1987). "The Interrelationships of Higher Ruminant Families with Special Emphasis on the Members of the Cervoidea". American Museum Novitates (2893): 1–85. hdl:2246/5180. Archived from the original on 6 October 2014. Retrieved 28 February 2016.
  28. 1 2 3 4 Janis, C.; Jarman, P. (1984). Macdonald, D. (ed.). The Encyclopedia of Mammals . New York: Facts on File. pp.  498–499. ISBN   978-0-87196-871-5.
  29. 1 2 3 Shively, C. L.; et al. (1985). "Some Aspects of the Nutritional Biology of the Collared Peccary". The Journal of Wildlife Management. 49 (3): 729–732. doi:10.2307/3801702. JSTOR   3801702.
  30. 1 2 3 "Artiodactyl". Encyclopædia Britannica Online. Encyclopædia Britannica, Inc. 2008. Retrieved 17 October 2008.
  31. Hewitt, David G (24 June 2011). Biology and Management of White-tailed Deer. ISBN   9781482295986.
  32. 1 2 Uwe Gille (2008). urinary and sexual apparatus, urogenital Apparatus. In: F.-V. Salomon and others (eds.): Anatomy for veterinary medicine. pp. 368–403. ISBN   978-3-8304-1075-1.
  33. Spinage, C. A. "Reproduction in the Uganda defassa waterbuck, Kobus defassa ugandae Neumann." Journal of reproduction and fertility 18.3 (1969): 445-457.
  34. Yong, Hwan-Yul. "Reproductive System of Giraffe (Giraffa camelopardalis)." Journal of Embryo Transfer 24.4 (2009): 293-295.
  35. Sumar, Julio. "Reproductive physiology in South American Camelids." Genetics of Reproduction in Sheep (2013): 81.
  36. 1 2 Pough, F. W.; Janis, C. M.; Heiser, J. B. (2005) [1979]. "Major Lineages of Mammals". Vertebrate Life (7th ed.). Pearson. p. 539. ISBN   978-0-13-127836-3.
  37. "Killer Whale". NOAA Fisheries. 3 August 2021. Retrieved 26 August 2021.
  38. McKie, Robin (22 September 2012). "Humans hunted for meat 2 million years ago". The Guardian. Retrieved 26 October 2015.
  39. "Bones From French Cave Show Neanderthals, Cro-Magnon Hunted Same Prey". ScienceDaily. 2003. Retrieved 17 October 2008.
  40. Clay, J. (2004). World Agriculture and the Environment: A Commodity-by-Commodity Guide to Impacts and Practices. Washington, D.C., US: Island Press. ISBN   978-1-55963-370-3.
  41. "Cetartiodactyla" . Retrieved 12 March 2007.
  42. "Artiodactyla". Encyclopedia of Life. Retrieved 15 November 2014.