Artiofabula is a

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

made up of the

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 ...

and the

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 S1 ...

. The clade was found in

molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

analyses and contradicted traditional relationships based on morphological analyses.

Etymology

The name Artiofabula was derived from Greek "artios" (meaning complete or perfect of its kind or, with respect to numbers, even), and Latin "fabula" (meaning fable). The latter referred to the clade breaking up traditional views on artiodactyl taxonomy based on morphological analyses, where camels grouped with ruminants, hippos with pigs, and whales were unrelated.

Phylogeny

Phylogenetic analyses of artiodactyls revealed the following relationships:(see e.g. Fig S10)

Classification

*Order

Artiodactyla Artiodactyls are placental mammals belonging to the order (biology), order Artiodactyla ( , ). Typically, they are ungulates which bear weight equally on two (an even number) of their five toes (the third and fourth, often in the form of a hoof ...

(even-toed ungulates) **Suborder

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 camel ...

(camelids) **Artiofabula (

ruminants Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principa ...

pigs The pig (''Sus domesticus''), also called swine (: swine) or hog, is an omnivorous, domesticated, even-toed, hoofed mammal. It is named the domestic pig when distinguishing it from other members of the genus '' Sus''. Some authorities cons ...

, peccaries,

whales Whales are a widely distributed and diverse group of fully Aquatic animal, aquatic placental mammal, placental marine mammals. As an informal and Colloquialism, colloquial grouping, they correspond to large members of the infraorder Cetacea ...

, and

dolphins A dolphin is an aquatic mammal in the cetacean clade Odontoceti (toothed whale). Dolphins belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontopori ...

) ***Suborder

(pigs and peccaries) **** Family

Suidae Suidae is a family (biology), family of Even-toed ungulate, artiodactyl mammals which are commonly called pigs, hogs, or swine. In addition to numerous fossil species, 18 Extant taxon, extant species are currently recognized (or 19 counting domes ...

19 species (pigs) **** Family Tayassuidae 4 species (peccaries) ***

(

, and

) ****Suborder

Ruminantia Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principa ...

(

antelope The term antelope refers to numerous extant or recently extinct species of the ruminant artiodactyl family Bovidae that are indigenous to most of Africa, India, the Middle East, Central Asia, and a small area of Eastern Europe. Antelopes do ...

, buffalo,

cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...

goats The goat or domestic goat (''Capra hircus'') is a species of goat-antelope that is mostly kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the famil ...

sheep Sheep (: sheep) or domestic sheep (''Ovis aries'') are a domesticated, ruminant mammal 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 d ...

deer A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) ...

giraffes 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. It is classified under the family Giraffidae, along with its closest extant re ...

, and

chevrotain Chevrotains, or mouse-deer, are small, even-toed ungulates that make up the family Tragulidae, and are the only living members of the infraorder Tragulina. The 10 extant species are placed in three genera, but several species also ar ...

s) *****Family

Antilocapridae The Antilocapridae are a family of ruminant artiodactyls endemic to North America. Their closest extant relatives are the giraffids. Only one species, the pronghorn (''Antilocapra americana''), is living today; all other members of the family ...

, 1 species (pronghorn) *****Family

Bovidae The Bovidae comprise the family (biology), biological family of cloven-hoofed, ruminant mammals that includes Bos, cattle, bison, Bubalina, buffalo, antelopes (including Caprinae, goat-antelopes), Ovis, sheep and Capra (genus), goats. A member o ...

, 135 species (

bison A bison (: bison) is a large bovine in the genus ''Bison'' (from Greek, meaning 'wild ox') within the tribe Bovini. Two extant taxon, extant and numerous extinction, extinct species are recognised. Of the two surviving species, the American ...

, buffalo,

, and

) *****Family

Cervidae A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) a ...

, 55 - 94 species (

elk The elk (: ''elk'' or ''elks''; ''Cervus canadensis'') or wapiti, is the second largest species within the deer family, Cervidae, and one of the largest terrestrial mammals in its native range of North America and Central and East Asia. ...

, and

moose The moose (: 'moose'; used in North America) or elk (: 'elk' or 'elks'; used in Eurasia) (''Alces alces'') is the world's tallest, largest and heaviest extant species of deer and the only species in the genus ''Alces''. It is also the tal ...

) *****Family

Giraffidae The Giraffidae are a family (biology), family of ruminant artiodactyl mammals that share a recent common ancestor with deer and bovids. This family, once a diverse group spread throughout Eurasia and Africa, presently comprises only two extant ge ...

, 2 species (

, okapis) *****Family

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 ...

, 4 - 7 species (musk deer) *****Family

Tragulidae Chevrotains, or mouse-deer, are small, even-toed ungulates that make up the family Tragulidae, and are the only living members of the infraorder Tragulina. The 10 extant species are placed in three genera, but several species also are ...

, 6 - 10 species (chevrotains, or mouse deer) ****Suborder

Whippomorpha Whippomorpha or Cetancodonta is a suborder of artiodactyls that contains all living cetaceans (whales, dolphins, and porpoises) and the hippopotamids. This makes it a crown group. Whippomorpha is a suborder within the order Artiodactyla (even-t ...

(aquatic or semi-aquatic even-toed ungulates) *****Infraorder

Ancodonta Ancodonta is an infraorder of 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 fossi ...

******Family

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 th ...

, 2 species (hippopotamuses) *****Infraorder

Cetacea Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...

(whales,

, and

porpoises Porpoises () are small dolphin-like cetaceans classified under the family Phocoenidae. Although similar in appearance to dolphins, they are more closely related to narwhals and belugas than to the true dolphins. There are eight extant speci ...

) ******Parvorder

Mysticeti Baleen whales (), also known as whalebone whales, are marine mammals of the parvorder Mysticeti in the infraorder Cetacea (whales, dolphins and porpoises), which use baleen plates (or "whalebone") in their mouths to sieve plankton from the wate ...

(baleen whales) *******Family

Balaenidae Balaenidae () is a Family (biology), family of whales of the parvorder Mysticeti (baleen whales) that contains mostly fossil taxa and two living genera: the right whale (genus ''Eubalaena''), and the closely related bowhead whale (genus ''Balaena ...

, 2 - 4 species (

right whales Right whales are three species of large baleen whales of the genus ''Eubalaena'': the North Atlantic right whale (''E. glacialis''), the North Pacific right whale (''E. japonica'') and the southern right whale (''E. australis''). They are class ...

and

bowhead whales The bowhead whale (''Balaena mysticetus''), sometimes called the Greenland right whale, Arctic whale, and polar whale, is a species of baleen whale belonging to the family Balaenidae and is the only living representative of the genus '' Balaena' ...

) *******Family

Balaenopteridae Rorquals () are the largest clade, group of baleen whales, comprising the family (biology), family Balaenopteridae, which contains nine extant taxon, extant species in two genus, genera. They include the largest known animal that has ever lived, ...

, 6 - 9 species ( rorquals) *******Family

Eschrichtiidae Eschrichtiidae or the gray whales is a family (taxonomy), family of baleen whale (Parvorder Mysticeti) with a single extant species, the gray whale (''Eschrichtius robustus''), as well as four described fossil genera: ''Archaeschrichtius'' (Mioce ...

, 1 species (

gray whale The gray whale (''Eschrichtius robustus''), also known as the grey whale,Britannica Micro.: v. IV, p. 693. is a baleen whale that migrates between feeding and breeding grounds yearly. It reaches a length of , a weight of up to and lives between ...

) *******Family Neobalaenidae, 1 species (

pygmy right whale The pygmy right whale (''Caperea marginata'') is a species of baleen whale. It may be a member of the cetotheres, a family of baleen whales which until 2012 were thought to be extinct; ''C. marginata'' has otherwise been considered the sole m ...

) ******Parvorder

Odontoceti The toothed whales (also called odontocetes, systematic name Odontoceti) are a parvorder of cetaceans that includes dolphins, porpoises, and all other whales with teeth, such as beaked whales and the sperm whales. 73 species of toothed whales ar ...

(toothed whales, dolphins, and porpoises) *******Superfamily Delphinoidea (

, arctic whales,

, and relatives) ********Family

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 ...

, 38 species (

, killer whales, and relatives) ********Family

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 Arcti ...

, 2 species (

beluga Beluga may refer to: Animals *Beluga (sturgeon) * Beluga whale Vehicles * Airbus Beluga, a large transport airplane * Airbus BelugaXL, a larger transport airplane * Beluga-class submarine, a class of Russian SSA diesel-electric submarine * U ...

and

narwhal The narwhal (''Monodon monoceros'') is a species of toothed whale native to the Arctic. It is the only member of the genus ''Monodon'' and one of two living representatives of the family Monodontidae. The narwhal is a stocky cetacean with a ...

) ********Family

Phocoenidae Porpoises () are small dolphin-like cetaceans classified under the family Phocoenidae. Although similar in appearance to dolphins, they are more closely related to narwhals and belugas than to the true dolphins. There are eight extant specie ...

, 6 species (

) *******Superfamily

Physeteroidea Physeteroidea is a superfamily (taxonomy), superfamily that includes three extant taxon, 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, t ...

(

sperm whales The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the genus '' Physeter'' and one of three extant species in the sperm whale super ...

) ********Family

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

, 2 species (

pygmy In anthropology, pygmy peoples are ethnic groups whose average height is unusually short. The term pygmyism is used to describe the phenotype of endemic short stature (as opposed to disproportionate dwarfism occurring in isolated cases in a po ...

and

dwarf sperm whale The dwarf sperm whale (''Kogia sima'') is a sperm whale that inhabits temperate and tropical oceans worldwide, in particular continental shelves and slopes. It was first described by biologist Richard Owen in 1866, based on illustrations by na ...

s) ********Family

Physeteridae 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 unit ...

, 1 species ( common sperm whale) *******Superfamily Ziphoidea ( beaked whales) ********Family Ziphidae, 22 species ( modern beaked whales) *******Superfamily

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 rive ...

(

river dolphins 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 rive ...

) ********Family

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 freshwater and marine deposits in the Neogene. The Amazon river dol ...

, 1 - 2 species ( Asian river dolphin(s)) *******Superfamily

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 rive ...

********Family

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 ...

, 1 - 3 species ( South American river dolphin(s)) ********Family

Pontoporiidae Pontoporiidae is a family of toothed whales, containing the extant genus '' Pontoporia'', and ten extinct genera. ''Pontoporia'' contains the sole extant species, the La Plata dolphin, which is endemic to South America South America is ...

, 1 species (

La Plata dolphin The La Plata dolphin, franciscana or toninha (''Pontoporia blainvillei'') is a species of river dolphin found in coastal Atlantic waters of southeastern South America. It is a member of the Inioidea group and the only one that lives in the ocean ...

) *******Superfamily

Lipotoidea Lipotidae is a Family (biology), 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'', w ...

********Family

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 ...

, 1 species (

baiji The baiji (''Lipotes vexillifer'') is a probably extinct species of freshwater dolphin native to the Yangtze river system in China. It is thought to be the first dolphin species driven to extinction due to the impact of humans. This dolphin is ...

or Chinese river dolphin)

References

{{Taxonbar, from=Q3428352 Artiodactyls Phylogenetics