Artiofabula is a

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, ...

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 in Eng ...

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 S10) ...

. The clade was found in molecular phylogenetic 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 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 ...

(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 camels are ...

(camelids) **Artiofabula (

ruminants 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 ...

pigs 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 ...

peccaries A peccary (also javelina or skunk pig) is a medium-sized, pig-like hoofed mammal of the family Tayassuidae (New World pigs). They are found throughout Central and South America, Trinidad in the Caribbean, and in the southwestern area of North A ...

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 ...

, 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 ...

) ***Suborder

(pigs and peccaries) **** Family

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 ...

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

(

, and

) ****Suborder

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 ...

(

antelope The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia. Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mammals ...

, buffalo,

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 ...

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

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 ...

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. Traditionally, giraffes were thought to be one species, ''Giraffa camelopardalis ...

, and

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

s) *****Family

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 ...

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

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, ...

, 135 species (

bison Bison are large bovines in the genus ''Bison'' (Greek: "wild ox" (bison)) within the tribe Bovini. Two extant and numerous extinct species are recognised. Of the two surviving species, the American bison, ''B. bison'', found only in North Ame ...

, buffalo,

, and

) *****Family

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 ...

, 55 - 94 species (

elk The elk (''Cervus canadensis''), also known as the wapiti, is one of the 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. The common ...

, and

moose The moose (in North America) or elk (in Eurasia) (''Alces alces'') is a member of the New World deer subfamily and is the only species in the genus ''Alces''. It is the largest and heaviest extant species in the deer family. Most adult mal ...

) *****Family

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 ...

, 2 species (

okapis The okapi (; ''Okapia johnstoni''), also known as the forest giraffe, Congolese giraffe, or zebra giraffe, is an artiodactyl mammal that is endemic to the northeast Democratic Republic of the Congo in central Africa. It is the only species ...

) *****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 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 v ...

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

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 ...

(aquatic or semi-aquatic even-toed ungulates) *****Infraorder Acodonta ******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 t ...

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

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 ...

(whales,

, 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 ...

) ******Parvorder

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 ...

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

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 ...

, 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'') is a species of baleen whale belonging to the family Balaenidae and the only living representative of the genus ''Balaena''. They are the only baleen whale endemic to the Arctic and subarctic waters, an ...

) *******Family

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 ...

, 6 - 9 species (

rorquals 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 whale ...

) *******Family

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

, 1 species (

gray whale The gray whale (''Eschrichtius robustus''), also known as the grey whale,Britannica Micro.: v. IV, p. 693. gray back whale, Pacific gray whale, Korean gray whale, or California gray whale, is a baleen whale that migrates between feeding and bree ...

) *******Family

Neobalaenidae Neobalaenidae is a family of baleen whales (suborder Mysticeti) including the extant pygmy right whale. Although traditionally considered related to balaenids, a recent phylogenetic study by Fordyce and Marx (2013) recovered the living pygmy ri ...

, 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 monotyp ...

) ******Parvorder

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 ...

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

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''., ...

(

, 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 The orca or killer whale (''Orcinus orca'') is a toothed whale belonging to the oceanic dolphin family, of which it is the largest member. It is the only extant species in the genus ''Orcinus'' and is recognizable by its black-and-white pa ...

, 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 Arctic ...

, 2 species (

beluga The beluga whale (/bɪˈluːɡə/) (Delphinapterus leucas) is an Arctic and sub-Arctic cetacean. It is one of two members of the family Monodontidae, along with the narwhal, and the only member of the genus Delphinapterus. It is also known as the wh ...

and

narwhal The narwhal, also known as a narwhale (''Monodon monoceros''), is a medium-sized toothed whale that possesses a large "tusk" from a protruding canine tooth. It lives year-round in the Arctic waters around Greenland, Canada and Russia. It is o ...

) ********Family

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 ...

, 6 species (

) *******Superfamily

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 ...

(

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 (biology), genus ''Physeter'' and one of three extant species in the s ...

) ********Family

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 ...

, 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 pop ...

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 n ...

s) ********Family Physeteridae, 1 species ( common sperm whale) *******Superfamily Ziphoidea (

beaked whales 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 ...

) ********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 riv ...

(

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 riv ...

) ********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 marine deposits in the Neogene. The Amazon river dolphin, Yangtze ...

, 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 riv ...

********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 a ...

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

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

) *******Superfamily

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 ...

********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 (; IPA: ; ''Lipotes vexillifer'', ''Lipotes'' meaning "left behind" and ''vexillifer'' "flag bearer") is a possibly extinct species of freshwater dolphin native to the Yangtze river system in China. It is thought to be the first dolph ...

or Chinese river dolphin)

References

{{Taxonbar, from=Q3428352 Even-toed ungulates Phylogenetics