Cetartiodactyla
Overview
Cetartiodactyla is the clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 in which whale
Whale
Whale is the common name for various marine mammals of the order Cetacea. The term whale sometimes refers to all cetaceans, but more often it excludes dolphins and porpoises, which belong to suborder Odontoceti . This suborder also includes the sperm whale, killer whale, pilot whale, and beluga...

s (including dolphin
Dolphin
Dolphins are marine mammals that are closely related to whales and porpoises. There are almost forty species of dolphin in 17 genera. They vary in size from and , up to and . They are found worldwide, mostly in the shallower seas of the continental shelves, and are carnivores, mostly eating...

s) and even-toed ungulate
Even-toed ungulate
The even-toed ungulates are ungulates whose weight is borne about equally by the third and fourth toes, rather than mostly or entirely by the third as in odd-toed ungulates such as horses....

s have currently been placed. The term was coined by merging the name for the two orders, Cetacea
Cetacea
The order Cetacea includes the marine mammals commonly known as whales, dolphins, and porpoises. Cetus is Latin and is used in biological names to mean "whale"; its original meaning, "large sea animal", was more general. It comes from Ancient Greek , meaning "whale" or "any huge fish or sea...

 and Artiodactyla, into a single word. The term Cetartiodactyla reflects the idea that whales evolved within the artiodactyls. Under this definition their closest living land relative is thought to be the hippopotamus
Hippopotamus
The hippopotamus , or hippo, from the ancient Greek for "river horse" , is a large, mostly herbivorous mammal in sub-Saharan Africa, and one of only two extant species in the family Hippopotamidae After the elephant and rhinoceros, the hippopotamus is the third largest land mammal and the heaviest...

. The clade formed by uniting whales and hippos is called Whippomorpha
Whippomorpha
Whippomorpha is a proposed clade containing the Cetacea and their closest living relatives, the hippopotamuses defined by Waddell et al. . It is defined as a node-based taxon, including all species that are descendants of the most recent common ancestor of Hippopotamus amphibius and Tursiops...

.
Encyclopedia
Cetartiodactyla is the clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 in which whale
Whale
Whale is the common name for various marine mammals of the order Cetacea. The term whale sometimes refers to all cetaceans, but more often it excludes dolphins and porpoises, which belong to suborder Odontoceti . This suborder also includes the sperm whale, killer whale, pilot whale, and beluga...

s (including dolphin
Dolphin
Dolphins are marine mammals that are closely related to whales and porpoises. There are almost forty species of dolphin in 17 genera. They vary in size from and , up to and . They are found worldwide, mostly in the shallower seas of the continental shelves, and are carnivores, mostly eating...

s) and even-toed ungulate
Even-toed ungulate
The even-toed ungulates are ungulates whose weight is borne about equally by the third and fourth toes, rather than mostly or entirely by the third as in odd-toed ungulates such as horses....

s have currently been placed. The term was coined by merging the name for the two orders, Cetacea
Cetacea
The order Cetacea includes the marine mammals commonly known as whales, dolphins, and porpoises. Cetus is Latin and is used in biological names to mean "whale"; its original meaning, "large sea animal", was more general. It comes from Ancient Greek , meaning "whale" or "any huge fish or sea...

 and Artiodactyla, into a single word. The term Cetartiodactyla reflects the idea that whales evolved within the artiodactyls. Under this definition their closest living land relative is thought to be the hippopotamus
Hippopotamus
The hippopotamus , or hippo, from the ancient Greek for "river horse" , is a large, mostly herbivorous mammal in sub-Saharan Africa, and one of only two extant species in the family Hippopotamidae After the elephant and rhinoceros, the hippopotamus is the third largest land mammal and the heaviest...

. The clade formed by uniting whales and hippos is called Whippomorpha
Whippomorpha
Whippomorpha is a proposed clade containing the Cetacea and their closest living relatives, the hippopotamuses defined by Waddell et al. . It is defined as a node-based taxon, including all species that are descendants of the most recent common ancestor of Hippopotamus amphibius and Tursiops...

. Alternatively, the term Cetartiodactyla is used to denote a clade where Cetacea evolved alongside Artiodactyla and not within it. Under this definition, all artiodactyls, including hippos, are more closely related to one another than any are to the whales.

This group has been proposed as a new order, but there is not sufficient evidence of the exact relationship between the two current orders to efficiently merge them.

Whales evolved from land mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s and appear to form a monophyletic group. It is well accepted that all whales evolved from a single ancestor. The most widely accepted hypothesis before the 1990s was that the closest relatives to whales were the fossil group Mesonychia. These were hooved
Hoof
A hoof , plural hooves or hoofs , is the tip of a toe of an ungulate mammal, strengthened by a thick horny covering. The hoof consists of a hard or rubbery sole, and a hard wall formed by a thick nail rolled around the tip of the toe. The weight of the animal is normally borne by both the sole...

, predominantly carnivorous, mammals that are known only from fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

s. But today, few authorities still consider mesonychids to be more closely related to whales than artiodactyls. Instead, they are usually considered to be the closest relative of the Cetartiodactyla as a whole.

Kin to hippos

The idea that whales evolved from within the Artiodactyla is based on analysis of DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...

 sequences. In the initial molecular analyses, whales were shown to be more closely related to ruminant
Ruminant
A ruminant is a mammal of the order Artiodactyla that digests plant-based food by initially softening it within the animal's first compartment of the stomach, principally through bacterial actions, then regurgitating the semi-digested mass, now known as cud, and chewing it again...

s (such as cattle
Cattle
Cattle are the most common type of large domesticated ungulates. They are a prominent modern member of the subfamily Bovinae, are the most widespread species of the genus Bos, and are most commonly classified collectively as Bos primigenius...

 and deer
Deer
Deer are the ruminant mammals forming the family Cervidae. Species in the Cervidae family include white-tailed deer, elk, moose, red deer, reindeer, fallow deer, roe deer and chital. Male deer of all species and female reindeer grow and shed new antlers each year...

) than ruminants are to pig
Pig
A pig is any of the animals in the genus Sus, within the Suidae family of even-toed ungulates. Pigs include the domestic pig, its ancestor the wild boar, and several other wild relatives...

s. In order for the order name to reflect a real evolutionary unit, the term Cetartiodactyla was coined.

Later molecular analyses included a wider sampling of artiodactyls and produced a more complete tale. Hippos were determined to be the closest relative of whales, ruminants were related to a whale/hippo clade, and pigs were more distant. In addition to producing the controversial whale/hippo clade, these analyses debunked the idea that hippos and pigs are closely related. This had been a popular taxonomic hypothesis (Suiformes) based on similarities in morphological
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

 (physical) characteristics.

In addition to DNA and protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

 sequences, researchers tracked the movement of transposon
Transposon
Transposable elements are sequences of DNA that can move or transpose themselves to new positions within the genome of a single cell. The mechanism of transposition can be either "copy and paste" or "cut and paste". Transposition can create phenotypically significant mutations and alter the cell's...

s called SINE
Siné
Maurice Sinet , known as Siné, is a French cartoonist.As a young man he studied drawing and graphic arts, while earning a living as a cabaret singer. After his military service he started publishing his drawings and also worked as a photo-retoucher for porn magazines. His first published drawing...

s in the genome
Genome
In modern molecular biology and genetics, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA. The genome includes both the genes and the non-coding sequences of the DNA/RNA....

 (see the method at retrotransposon marker
Retrotransposon Marker
Retrotransposon markers are retrotransposons that are used as cladistic markers.The analysis of SINEs – Short INterspersed Elements – LINEs – Long INterspersed Elements – or truncated LTRs – Long Terminal Repeats – as molecular cladistic markers represents a particularly interesting complement to...

). A transposon is a DNA sequence that will occasionally make a copy of itself and insert that copy into another part of the genome. It is considered highly unlikely that SINEs will insert themselves into exactly the same part of a genome by chance. The data indicate that several transposons inserted themselves at the same point in the genomes of whales, ruminants and hippos (sometimes referred to as "pseudoruminants" because although they have four-chambered stomachs like true ruminants, they do not chew the cud). This insertion point is not shared with camel
Camel
A camel is an even-toed ungulate within the genus Camelus, bearing distinctive fatty deposits known as humps on its back. There are two species of camels: the dromedary or Arabian camel has a single hump, and the bactrian has two humps. Dromedaries are native to the dry desert areas of West Asia,...

s and pigs.

This hypothesis has been tested with DNA sequences from a host of genes: the complete mitochondrial genome (as well as several of its genes independently), beta-casein, kappa-casein, von Willebrand factor
Von Willebrand factor
von Willebrand factor is a blood glycoprotein involved in hemostasis. It is deficient or defective in von Willebrand disease and is involved in a large number of other diseases, including thrombotic thrombocytopenic purpura, Heyde's syndrome, and possibly hemolytic-uremic syndrome...

, breast cancer 1, recombination activating gene
Recombination activating gene
The recombination activating genes encode enzymes that play an important role in the rearrangement and recombination of the genes of immunoglobulin and T cell receptor molecules during the process of VDJ recombination...

s 1 and 2, cannabinoid receptor
Cannabinoid receptor
The cannabinoid receptors are a class of cell membrane receptors under the G protein-coupled receptor superfamily. As is typical of G protein-coupled receptors, the cannabinoid receptors contain seven transmembrane spanning domains...

 1, and several others. These sequence data and the transposons converge on the same conclusion that hippos and whales are more closely related to one another than either is to other artiodactyls.

Sequences analyzed in combined analyses with morphological characters have also produced the same results as sequences alone. Some have argued that the sheer number of characters (one for each nucleotide
Nucleotide
Nucleotides are molecules that, when joined together, make up the structural units of RNA and DNA. In addition, nucleotides participate in cellular signaling , and are incorporated into important cofactors of enzymatic reactions...

) in sequences swamps out the effects of morphology. There have been a few morphology-based studies that have suggested (weakly) the same results as the molecular results, but overall most morphological studies have conflicted with the whale/hippo hypothesis of Cetartiodactyla.

An important exception is a recent study conducted by Boisserie et al. (2005). They examined 80 hard morphological characters of fossil and extant cetartiodactylan taxa. Their results suggest that hippopotamids evolved from within a clade of anthracotheres. That anthracothere/hippopotamid clade appears to be sister to the Cetacea and supports the molecular results.

A clade apart

The morphology and paleontology
Paleontology
Paleontology "old, ancient", ὄν, ὀντ- "being, creature", and λόγος "speech, thought") is the study of prehistoric life. It includes the study of fossils to determine organisms' evolution and interactions with each other and their environments...

-based opinion of the relationship of whales to other mammals has also undergone considerable changes since the early 1990s. Fossils such as Rodhocetus
Rodhocetus
Rodhocetus is one of several extinct whale genera that possess land mammal characteristics, thus demonstrating the evolutionary transition from land to sea.-Description:...

, have been discovered that refute the notion that whales are derived from or are closely related to the mesonychid
Mesonychid
Mesonychia are an extinct order of medium to large-sized carnivorous mammals that were closely related to artiodactyls and to cetaceans...

s. Many morphologists and paleontologists support the notion of a clade called Cetartiodactyla that unites Cetacea + Artiodactyla. Many are not, however, in support of the hypothesis that Cetacea evolved from within the Artiodactyla. Under this definition, Artiodactyla remains a valid clade since artiodactyls evolved from a common ancestor separate from whales. Cetartiodactyla would represent a grandorder or superorder uniting the two orders.

The vast majority of phylogenetic analyses based on morphological characters have not uncovered a whale/hippo clade, but show Cetacea and Artiodactyla as distinct from one another. Features of the bones of the astragalus
Talus bone
-External links:* *...

 in the ankle region are cited as particular evidence for a monophyletic Artiodactyla. In any case, an origin of the Cetacea from within the Ungulata was proposed as early as 1900 by Frank Evers Beddard
Frank Evers Beddard
Frank Evers Beddard FRS was a British zoologist. He won the Linnean Medal in 1916 for his book on oligochaetes.Beddard was born in Dudley, educated at Harrow and studied at New College, Oxford. He was naturalist to the Challenger Expedition Commission from 1882 to 1884...

 in A book of whales. But Beddard found the anatomical and fossil record at that time insufficient to rule out other possibilities, such as the notion that whales are a basal lineage among placental mammals, or the popular theory relating them to terrestrial carnivores. Because Beddard himself was not fully convinced, his bold proposal was largely forgotten.

Hippo fossils are not observed until the Miocene
Miocene
The Miocene is a geological epoch of the Neogene Period and extends from about . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the Pliocene. The Miocene follows the Oligocene...

, but whale ancestors have been dated to the Eocene
Eocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...

. The whale/hippo hypothesis imposes a gap of nearly 30 million years where no hippo ancestors are present. Certain anthracotheres have been proposed as hippo ancestors, but this concept has not garnered wide support among paleontologists. However, the most recent hypothesis into the origins of hippopotamids suggests that hippos and whales shared a common semi-aquatic ancestor that branched off from other Artiodactyls around . This hypothesized ancestral group likely split into two branches around . One branch would evolve into cetaceans
Evolution of cetaceans
The cetaceans are marine mammal descendants of land mammals. Their terrestrial origins are indicated by:* Their need to breathe air from the surface;* The bones of their fins, which resemble the limbs of land mammals...

, possibly beginning with the proto-whale Pakicetus
Pakicetus
Pakicetus is a genus of extinct terrestrial carnivorous mammal of the family Pakicetidae which was endemic to Pakistan from the Eocene .Pakicetus existed for approximately...

from with other early whale ancestors collectively known as Archaeoceti
Archaeoceti
Archaeocetes, or "ancient whales", are a paraphyletic group of cetaceans that gave rise to the modern cetaceans.The archaeocetes were once thought to have evolved from the mesonychids, based on dental characteristics...

, which eventually underwent aquatic adaptation
Aquatic adaptation
Several animal groups have undergone aquatic adaptation, going from being purely terrestrial animals to living at least part of the time in water. The adaptations in early speciation tend to develop as the animal ventures into water in order to find available food. As successive generations spend...

 into the completely aquatic cetacea
Cetacea
The order Cetacea includes the marine mammals commonly known as whales, dolphins, and porpoises. Cetus is Latin and is used in biological names to mean "whale"; its original meaning, "large sea animal", was more general. It comes from Ancient Greek , meaning "whale" or "any huge fish or sea...

ns. Thus, Cetartiodactyla is supported as a monophyletic group.
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