Ungulates ( ) are members of the diverse

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

Ungulata which primarily consists of large mammals with

hooves The hoof (plural: hooves) is the tip of a toe of an ungulate mammal, which is covered and strengthened with a thick and horny keratin covering. Artiodactyls are even-toed ungulates, species whose feet have an even number of digits, yet the rumi ...

. These include

odd-toed ungulate Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) ...

s such as

horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million y ...

rhinoceros A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct species o ...

es, and

tapir Tapirs ( ) are large, herbivorous mammals belonging to the family Tapiridae. They are similar in shape to a pig, with a short, prehensile nose trunk. Tapirs inhabit jungle and forest regions of South and Central America, with one species inhabit ...

s; and

even-toed ungulate The even-toed ungulates (Artiodactyla , ) 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 poster ...

s such as

cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult mal ...

pig The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus s ...

giraffe The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, ''Giraffa camelopardalis ...

camel A camel (from: la, camelus and grc-gre, κάμηλος (''kamēlos'') from Hebrew or Phoenician: גָמָל ''gāmāl''.) is an even-toed ungulate in the genus ''Camelus'' that bears distinctive fatty deposits known as "humps" on its back. C ...

sheep Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus ''Ovis'', in everyday usage it almost always refers to domesticated s ...

deer Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer ...

, and

hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extan ...

es.

Cetacean Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...

s such as

whales Whales are a widely distributed and diverse group of fully aquatic placental marine mammals. As an informal and colloquial grouping, they correspond to large members of the infraorder Cetacea, i.e. all cetaceans apart from dolphins and ...

dolphins A dolphin is an aquatic mammal within the infraorder Cetacea. Dolphin species belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontoporiidae (t ...

, and

porpoises Porpoises are a group of fully aquatic marine mammals, all of which are classified under the family Phocoenidae, parvorder Odontoceti (toothed whales). Although similar in appearance to dolphins, they are more closely related to narwhals an ...

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. The term means, roughly, "being hoofed" or "hoofed animal". As a descriptive term, "ungulate" normally excludes cetaceans as they do not possess most of the typical morphological characteristics of other ungulates, but recent discoveries indicate that they were also descended from early

artiodactyls The even-toed ungulates (Artiodactyla , ) 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 poster ...

. Ungulates are typically herbivorous and many employ specialized

gut bacteria Gut microbiota, gut microbiome, or gut flora, are the microorganisms, including bacteria, archaea, fungi, and viruses that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the gut mic ...

to allow them to digest cellulose. Some modern species, such as

s, are omnivorous, while some prehistoric species, such as

mesonychia Mesonychia ("middle claws") 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 ...

ns, were carnivorous.

Etymology

Ungulate is from the

Late Latin Late Latin ( la, Latinitas serior) is the scholarly name for the form of Literary Latin of late antiquity.Roberts (1996), p. 537. English dictionary definitions of Late Latin date this period from the , and continuing into the 7th century in t ...

adjective ''ungulatus'', "hoofed". ''Ungulatus'' is a

diminutive A diminutive is a root word that has been modified to convey a slighter degree of its root meaning, either to convey the smallness of the object or quality named, or to convey a sense of intimacy or endearment. A (abbreviated ) is a word-formati ...

form of Latin ''unguis'', "nail" (finger nail; toe nail).

Classifications

History

Ungulata is a clade (or in some taxonomies, a grand order) of mammals. The two orders of ungulates were the Perissodactyla (odd-toed ungulates) and Artiodactyla (even-toed ungulates). Hyracoidea (hyraxes), Sirenia (sea cows) (dugongs and manatees) and Proboscidea (elephants) were in the past included in a superorder called Paenungulata which was grouped with the ungulata. These three orders were now considered a clade and grouped in the Afrotheria clade while Ungulata is now grouped under the

Laurasiatheria Laurasiatheria ("laurasian beasts") is a superorder of placental mammals that groups together true insectivores ( eulipotyphlans), bats ( chiropterans), carnivorans, pangolins ( pholidotes), even-toed ungulates (artiodactyls), odd-toed ungulates ...

clade. In 2009 morphological and molecular work found that aardvarks, hyraxes, sea cows, and elephants were more closely related to each other and to

sengi Elephant shrews, also called jumping shrews or sengis, are small insectivorous mammals native to Africa, belonging to the family Macroscelididae, in the order Macroscelidea. Their traditional common English name "elephant shrew" comes from a pe ...

tenrec A tenrec is any species of mammal within the afrotherian family Tenrecidae endemic to Madagascar. Tenrecs are wildly diverse; as a result of convergent evolution some resemble hedgehogs, shrews, opossums, rats, and mice. They occupy aquatic, a ...

s, and

golden mole Golden moles are small insectivorous burrowing mammals endemic to Sub-Saharan Africa. They comprise the family Chrysochloridae and as such they are taxonomically distinct from the true moles, family Talpidae, and other mole-like families, all o ...

s than to the perissodactyls and artiodactyls, and form the clade

Afrotheria Afrotheria ( from Latin ''Afro-'' "of Africa" + ''theria'' "wild beast") is a clade of mammals, the living members of which belong to groups that are either currently living in Africa or of African origin: golden moles, elephant shrews (also know ...

. Elephants, sea cows, and hyraxes were grouped together in the clade

Paenungulata Paenungulata (from Latin ''paene'' "almost" + ''ungulātus'' "having Hoof, hoofs") is a clade of "sub-Ungulate, ungulates", which groups three Extant taxon, extant mammal Order (biology), orders: Proboscidea (including elephants), Sirenia (sirenia ...

, while the aardvark has been considered as either a close relative to them or a close relative to sengis in the clade

Afroinsectiphilia The Afroinsectiphilia (African insectivores) is a clade that has been proposed based on the results of recent molecular phylogenetic studies. Many of the taxa within it were once regarded as part of the order Insectivora, but Insectivora is now ...

. This is a striking example of

convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...

. There is now some dispute as to whether this smaller Ungulata is a

cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...

(evolution-based) group, or merely a

phenetic In biology, phenetics ( el, phainein – to appear) , also known as taximetrics, is an attempt to classify organisms based on overall similarity, usually in morphology or other observable traits, regardless of their phylogeny or evolutionary rel ...

group (

form taxon Form classification is the classification of organisms based on their morphology, which does not necessarily reflect their biological relationships. Form classification, generally restricted to palaeontology, reflects uncertainty; the goal of sc ...

) or

folk taxon A folk taxonomy is a vernacular name, naming system, as distinct from Taxonomy (general), scientific taxonomy. Folk biological classification is the way people traditionally describe and organize their natural surroundings/the world around them, ...

(similar, but not necessarily related). Some studies have indeed found the

mesaxonia Mesaxonia (near-synonymous with Panperissodactyla) is a clade of ungulates whose weight is distributed on the third toe on all legs through the plane symmetry of their feet. For a while it was often seen to only contain the order Perissodactyla ...

n ungulates and paraxonian ungulates to form a monophyletic lineage, closely related to either the

Ferae Ferae ( , , "wild beasts") is a mirorder of placental mammalsMalcolm C. McKenna, Susan K. Bell: ''Classification of Mammals: Above the Species Level'' in Columbia University Press, New York (1997), 631 Seiten. that groups together clades Pan-Ca ...

(the

carnivora Carnivora is a Clade, monophyletic order of Placentalia, placental mammals consisting of the most recent common ancestor of all felidae, cat-like and canidae, dog-like animals, and all descendants of that ancestor. Members of this group are f ...

ns and the

pangolin Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (, from Ancient Greek ϕολιδωτός – "clad in scales"). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smut ...

s) in the clade

Fereuungulata Ferungulata ("wild beasts and ungulates") is a grandorder of placental mammals that groups together mirorder Ferae and clade Pan-Euungulata. It has existed in two guises, a traditional one based on morphological analysis and a revised one taki ...

or to the

bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...

s. Other studies found the two orders not that closely related, as some place the perissodactyls as close relatives to bats and Ferae in

Pegasoferae Pegasoferae ("winged beast") is a proposed clade of mammals based on genomic research in molecular systematics by Nishihara, Hasegawa and Okada (2006). To the surprise of the authors, their data led them to propose a clade that includes bats (or ...

and others place the artiodactyls as close relatives to bats.

Taxonomy

Below is a simplified taxonomy (assuming that ungulates do indeed form a natural grouping) with the extant families, in order of the relationships. Keep in mind that there were still some grey areas of conflict, such as the case with relationship of the

pecora 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 “Pecor ...

n families and the

baleen whale Baleen whales (systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea (whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in their ...

families. See each family for the relationships of the species as well as the controversies in their respective article. * Ungulata (= Euungulata) ** Perissodactyla (Mesaxonian ungulates) *** Hippomorpha ****

Equidae Equidae (sometimes known as the horse family) is the taxonomic family of horses and related animals, including the extant horses, asses, and zebras, and many other species known only from fossils. All extant species are in the genus '' Equus'', ...

: Horses, asses and zebras ***

Ceratomorpha Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) ...

****

Tapiridae Tapirs ( ) are large, herbivorous mammals belonging to the family Tapiridae. They are similar in shape to a pig, with a short, prehensile nose trunk. Tapirs inhabit jungle and forest regions of South and Central America, with one species inhabi ...

: Tapirs ****

Rhinocerotidae A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct species ...

: Rhinoceroses **

Artiodactyla The even-toed ungulates (Artiodactyla , ) 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 poster ...

Cetartiodactyla The even-toed ungulates (Artiodactyla , ) 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 poster ...

) (Paraxonian ungulates) ***

Tylopoda Tylopoda (meaning "calloused foot") 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 ...

****

Camelidae Camelids are members of the biological family Camelidae, the only currently living family in the suborder Tylopoda. The seven extant members of this group are: dromedary camels, Bactrian camels, wild Bactrian camels, llamas, alpacas, vicuñas, ...

: Camels and llamas *** Artiofabula ****

Suina Suina (also known as Suiformes) is a suborder of omnivorous, non-ruminant artiodactyl mammals that includes the domestic pig and peccaries. A member of this clade is known as a suine. Suina includes the family Suidae, termed suids, known in Eng ...

***** Tayassuidae: Peccaries *****

Suidae Suidae is a family of artiodactyl mammals which are commonly called pigs, hogs or swine. In addition to numerous fossil species, 18 extant species are currently recognized (or 19 counting domestic pigs and wild boars separately), classified into ...

: Pigs ****

Cetruminantia The Cetruminantia are a clade made up of the Cetancodontamorpha (or Whippomorpha) and their closest living relatives, the Ruminantia. Cetruminantia's placement within Artiodactyla can be represented in the following cladogram: (see fig S10) ...

*****

Ruminantia Ruminants ( suborder Ruminantia) are hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microbial actions. The ...

******

Tragulidae The asterisk ( ), from Late Latin , from Ancient Greek , ''asteriskos'', "little star", is a typographical symbol. It is so called because it resembles a conventional image of a heraldic star. Computer scientists and mathematicians often voc ...

: Chevrotains ****** Cervoidea *******

Antilocapridae The Antilocapridae are a family of artiodactyls endemic to North America. Their closest extant relatives are the giraffids with which they comprise the superfamily Giraffoidea. Only one species, the pronghorn (''Antilocapra americana''), is li ...

: Pronghorn *******

Giraffidae The Giraffidae are a family of ruminant artiodactyl mammals that share a common ancestor with deer and bovids. This family, once a diverse group spread throughout Eurasia and Africa, presently comprises only two extant genera, the giraffe (one or ...

: Giraffes and okapi *******

Cervidae Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer ...

: Deer *******

Moschidae Moschidae is a family of pecoran even-toed ungulates, containing the musk deer (''Moschus'') and its extinct relatives. They are characterized by long 'saber teeth' instead of horns, antlers or ossicones, modest size (''Moschus'' only reaches ; ...

: Musk deer *******

Bovidae The Bovidae comprise the biological family of cloven-hoofed, ruminant mammals that includes cattle, bison, buffalo, antelopes, and caprines. A member of this family is called a bovid. With 143 extant species and 300 known extinct species, ...

: Cattle and antelopes *****

Whippomorpha Whippomorpha or Cetancodonta is a group of animals that contains all living cetaceans (whales, dolphins, etc.) and hippopotamuses, as well as their extinct relatives, i.e. Entelodont, Entelodonts and Andrewsarchus. All Whippomorphs are descendan ...

******

Hippopotamidae 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 t ...

: Hippopotamuses ******

Cetacea Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...

*******

Mysticeti Baleen whales (systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea (whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in their ...

********

Balaenidae Balaenidae () is a family of whales of the parvorder Mysticeti that contains two living genera: the right whales (genus ''Eubalaena''), and in a separate genus, the closely related bowhead whale (genus ''Balaena''). Evolutionary history Bale ...

: Bowhead and right whales ********

Cetotheriidae Cetotheriidae is a family of baleen whales (parvorder Mysticeti). The family is known to have existed from the Late Oligocene to the Early Pleistocene before going extinct. Although some phylogenetic studies conducted by recovered the living py ...

: Pygmy right whale ********

Balaenopteridae Rorquals () are the largest group of baleen whales, which comprise the family Balaenopteridae, containing ten extant species in three genera. They include the largest animal that has ever lived, the blue whale, which can reach , and the fin wha ...

: Rorquals *******

Odontoceti The toothed whales (also called odontocetes, systematic name Odontoceti) are a parvorder of cetaceans that includes dolphins, porpoises, and all other whales possessing teeth, such as the beaked whales and sperm whales. Seventy-three species of ...

********

Physeteroidea Physeteroidea is a superfamily that includes three extant species of whales: the sperm whale, in the genus ''Physeter'', and the pygmy sperm whale and dwarf sperm whale, in the genus ''Kogia''. In the past, these genera have sometimes been united ...

********* Physeteridae: Sperm whale *********

Kogiidae Kogiidae is a family comprising at least two extant species of Cetacea, the Pygmy sperm whale, pygmy (''Kogia breviceps)'' and dwarf sperm whale, dwarf (''K. sima)'' sperm whales. As their common names suggest, they somewhat resemble sperm whales ...

: Lesser sperm whales ********

Platanistoidea River dolphins are a polyphyletic group of fully aquatic mammals that reside exclusively in freshwater or brackish water. They are an informal grouping of dolphins, which itself is a paraphyletic group within the infraorder Cetacea. Extant riv ...

*********

Platanistidae Platanistidae is a family of river dolphins containing the extant Ganges river dolphin and Indus river dolphin (both in the genus '' Platanista'') but also extinct relatives from marine deposits in the Neogene. The Amazon river dolphin, Yangtze ...

: Indian river dolphins ********

Ziphioidea Beaked whales (systematic name Ziphiidae) are a family of cetaceans noted as being one of the least known groups of mammals because of their deep-sea habitat and apparent low abundance. Only three or four of the 24 species are reasonably well-k ...

*********

Ziphiidae Beaked whales (systematic name Ziphiidae) are a family of cetaceans noted as being one of the least known groups of mammals because of their deep-sea habitat and apparent low abundance. Only three or four of the 24 species are reasonably well-kn ...

: Beaked whales ********

Lipotoidea Lipotidae is a family of river dolphins containing the possibly extinct baiji of China and the fossil genus ''Parapontoporia'' from the Late Miocene and Pliocene of the Pacific coast of North America. The genus '' Prolipotes'', which is based on ...

*********

Lipotidae Lipotidae is a family of river dolphins containing the possibly extinct baiji of China and the fossil genus ''Parapontoporia'' from the Late Miocene and Pliocene of the Pacific coast of North America. The genus '' Prolipotes'', which is based on ...

: Baiji (functionally extinct) ********

Inioidea River dolphins are a polyphyletic group of fully aquatic mammals that reside exclusively in freshwater or brackish water. They are an informal grouping of dolphins, which itself is a paraphyletic group within the infraorder Cetacea. Extant riv ...

*********

Iniidae Iniidae is a family of river dolphins containing one living genus, ''Inia'', and four extinct genera. The extant genus inhabits the river basins of South America, but the family formerly had a wider presence across the Atlantic Ocean. Iniidae a ...

: Amazonian river dolphins ********* Pontoporiidae: La Plata dolphin ********

Delphinoidea Delphinoidea is the largest group of toothed whales with 66 genera in 6 families. The largest living member of the superfamily is the killer whale, which can reach 6 tons, while the smallest is the vaquita. Taxonomy Based on McGowen ''et al''., ...

*********

Monodontidae The cetacean family Monodontidae comprises two living whale species, the narwhal and the beluga whale and at least four extinct species, known from the fossil record. Beluga and Narwhal are native to coastal regions and pack ice around the Arctic ...

: Beluga and narwhal *********

Phocoenidae Porpoises are a group of fully aquatic marine mammals, all of which are classified under the family Phocoenidae, parvorder Odontoceti (toothed whales). Although similar in appearance to dolphins, they are more closely related to narwhals an ...

: Porpoises *********

Delphinidae Oceanic dolphins or Delphinidae are a widely distributed family of dolphins that live in the sea. Close to forty extant species are recognised. They include several big species whose common names contain "whale" rather than "dolphin", such as the ...

: Oceanic dolphins

Phylogeny

Below is the general consensus of the phylogeny of the ungulate families.

Evolutionary history

Cladogram of Cetacea within Artiodactyla

Perissodactyla and

include the majority of large land mammals. These two groups first appeared during the late

Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...

, rapidly spreading to a wide variety of species on numerous continents, and have developed in parallel since that time. Some scientists believed that modern ungulates were descended from an

evolutionary grade A grade is a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit. Definition An evolutionary grade is a group of ...

of mammals known as the

condylarth Condylarthra is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene epochs. They are considered early, primitive ungulates. It is now largely considered to be a wast ...

s; the earliest known member of the group was the tiny ''

Protungulatum ''Pro