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Temporal range: Late Jurassic or Early CretaceousHolocene, 161 or 145–0 Ma
Juramaia NT.jpg
Juramaia , the oldest known eutherian
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Subclass: Theria
Clade: Eutheria
Gill, 1872

Eutheria ( /jˈθɪəriə/ ; from Greek εὐ-, eu- "good" or "right" and θηρίον, thēríon "beast" hence "true beasts") is one of two mammalian clades with extant members that diverged in the Early Cretaceous or perhaps the Late Jurassic. All placental mammals indigenous to Europe, Africa, Asia, the Americas, Australia, and Antarctica are eutherians. Extant eutherians, their last common ancestor, and all extinct descendants of that ancestor are members of Placentalia.


Eutherians are distinguished from noneutherians by various phenotypic traits of the feet, ankles, jaws and teeth. All extant eutherians lack epipubic bones, which are present in all other living mammals (marsupials and monotremes). This allows for expansion of the abdomen during pregnancy. [2]

The oldest-known eutherian species is Juramaia sinensis , dated at 161  million years ago from the Jurassic in China. [3]

Eutheria was named in 1872 by Theodore Gill; in 1880 Thomas Henry Huxley defined it to encompass a more broadly defined group than Placentalia. [4]


The entocuneiform bone Cambridge Natural History Mammalia Fig 068.png
The entocuneiform bone

Distinguishing features are:

Evolutionary history

Eutheria contains several extinct genera as well as larger groups, many with complicated taxonomic histories still not fully understood. Members of the Adapisoriculidae, Cimolesta and Leptictida have been previously placed within the out-dated placental group Insectivora, while Zhelestids have been considered primitive ungulates. [6] However, more recent studies have suggested these enigmatic taxa represent stem group eutherians, more basal to Placentalia. [7] [8]

The weakly favoured cladogram favours Boreoeuthearia as a basal eutherian clade as sister to the Atlantogenata. [9] [10] [11]







The fossil eutherian species believed to be the oldest known is Juramaia sinensis , which lived about 160  million years ago. [3] Montanalestes was found in North America, while all other nonplacental eutherian fossils have been found in Asia. The earliest-known placental fossils have also been found in Asia. [5]



Other mammaliaformes








Simplified, non-systematic, outline of evolution of eutheria from cynodont therapsids. [5]
† = extinct

Related Research Articles

Marsupial Infraclass of mammals in the clade Metatheria

Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a pouch. Well-known marsupials include kangaroos, wallabies, koalas, opossums, wombats, Tasmanian devils, and the extinct thylacine. Some lesser-known marsupials are the dunnarts, potoroos, and the cuscus.

Placentalia Infraclass of mammals in the clade Eutheria

Placentalia is one of the three extant subdivisions of the class of animals Mammalia; the other two are Monotremata and Marsupialia. The placentals are partly distinguished from other mammals in that the fetus is carried in the uterus of its mother to a relatively late stage of development. The name is something of a misnomer considering that marsupials also nourish their fetuses via a placenta, though for a relatively briefer period, giving birth to less developed young who are then kept for a period in the mother's pouch.


Eomaia is a genus of extinct fossil mammals containing the single species Eomaia scansoria, discovered in rocks that were found in the Yixian Formation, Liaoning Province, China, and dated to the Barremian Age of the Lower Cretaceous about 125 million years ago. The single fossil specimen of this species is 10 centimetres (3.9 in) in length and virtually complete. An estimate of the body weight is between 20–25 grams (0.71–0.88 oz). It is exceptionally well-preserved for a 125-million-year-old specimen. Although the fossil's skull is squashed flat, its teeth, tiny foot bones, cartilages and even its fur are visible.


Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a slightly more inclusive group than the marsupials; it contains all marsupials as well as many extinct non-marsupial relatives.

Zalambdalestes Genus of shrew-like mammal from the Upper Cretaceous period

Zalambdalestes was a eutherian mammal, most likely not a placental due to the presence of an epipubic bone, living during the Upper Cretaceous in Mongolia.


Pucadelphys is an extinct genus of non-marsupial metatherian species. The genus contains a single species, P. andinus. Fossils of Pucadelphys have been found in the Santa Lucía Formation in Tiupampa in Bolivia.


Sinodelphys is an extinct mammal from the Early Cretaceous. To date, it is the oldest metatherian fossil known, estimated to be 125 million years old. It was discovered and described in 2003 in rocks of the Yixian Formation in Liaoning Province, China, by a team of scientists including Zhe-Xi Luo and John Wible.


Protungulatum is an extinct genus of mammal first found in the Bug Creek Anthills in northeastern Montana. The Bug Creek Anthills were initially believed to be Late Cretaceous because of the presence of the remains of non-avian dinosaurs and common Cretaceous mammals, but these were later shown to have been reworked from Late Cretaceous strata, and consequently the Bug Creek Anthills are currently believed to be Early Paleocene (Puercan) in age. Remains from the Ravenscrag Formation of Saskatchewan, Canada have been assigned to P. donnae. These remains may also be Cretaceous in age, but the age of the Ravenscrag Formation is not entirely certain. In 2011, remains of a new species of Protungulatum, P. coombsi, from the Hell Creek Formation, which is definitely Late Cretaceous (Maastrichtian) in age, proved that Protungulatum was present in both the Cretaceous and the Paleocene. It was initially assigned to the order condylarthra, a group of archaic "ungulates", that is now known to be polyphyletic. According to Archibald et al. (2011), Protungulatum is not even definitely a placental mammal. Some studies have found it to be close to Cetartiodactyla, but the most recent analysis holds it to be a non-placental eutherian.

Cimolestes is a genus of early eutherians with a full complement of teeth adapted for eating insects and other small animals. Paleontologists have disagreed on its relationship to other mammals, in part because quite different animals were assigned to the genus, making Cimolestes a grade taxon of animals with similar features rather than a genus of closely related ones. Fossils have been found in North America, where they first appeared during the Late Cretaceous. According to some paleontologists, they died out at the start of the Paleocene, while others report the genus from the early Eocene.

Evolution of mammals Derivation of mammals from a synapsid precursor, and the adaptive radiation of mammal species

The evolution of mammals has passed through many stages since the first appearance of their synapsid ancestors in the Pennsylvanian sub-period of the late Carboniferous period. By the mid-Triassic, there were many synapsid species that looked like mammals. The lineage leading to today's mammals split up in the Jurassic; synapsids from this period include Dryolestes, more closely related to extant placentals and marsupials than to monotremes, as well as Ambondro, more closely related to monotremes. Later on, the eutherian and metatherian lineages separated; the metatherians are the animals more closely related to the marsupials, while the eutherians are those more closely related to the placentals. Since Juramaia, the earliest known eutherian, lived 160 million years ago in the Jurassic, this divergence must have occurred in the same period.

Maelestes is a prehistoric shrew-like mammal discovered in 1997 in the Gobi Desert. The animal lived in the late Cretaceous Period, around 71-75 million years ago, and was a contemporary of dinosaurs such as Velociraptor and Oviraptor. According to some scientists, the discovery and analysis of this species suggests that true placental mammals appeared near the time the non-avian dinosaurs became extinct 66 million years ago, not much earlier in the Cretaceous as thought by others. However, the presence of an epipubic bone, among other characteristics, place it as a non-placental eutherian.

Epipubic bone

Epipubic bones are a pair of bones projecting forward from the pelvic bones of modern marsupials and most non-placental fossil mammals: multituberculates, monotremes, and even basal eutherians. They first occur in non-mammalian cynodonts such as tritylodontids, suggesting that they are a synapomorphy between them and Mammaliformes.


Vincelestes is an extinct genus of actively mobile mammal, that lived in what would be South America during the Early Cretaceous from 130—112 mya, existing for approximately 18 million years .

Dryolestida is an extinct order of mammals; most of the members are mostly known from the Jurassic to Paleogene, with one member, Necrolestes, surviving as late as the early Miocene. It has been suggested that these mammals are either the possible ancestors of therian mammals or an offshoot from the same evolutionary line. It is also believed that they developed a fully mammalian jaw and also had the three middle ear bones. Other than that, not much is known about them, this is because their fossils are made up mostly of jaw and tooth remains.


Leptictida is a possibly invalid extinct order of placental mammals. Their classification is contentious: according to cladistic studies, they may be (distantly) related to Euarchontoglires, although they are more recently regarded as the first branch to split from basal eutherians. One recent large-scale cladistic analysis of eutherian mammals favored lepictidans as close to the placental crown-clade; and several other recent analyses that included data from Cretaceous non-eutherian mammals found Leptictis to belong to the superorder Afrotheria.

Several mammals are known from the Mesozoic of Madagascar. The Bathonian Ambondro, known from a piece of jaw with three teeth, is the earliest known mammal with molars showing the modern, tribosphenic pattern that is characteristic of marsupial and placental mammals. Interpretations of its affinities have differed; one proposal places it in a group known as Australosphenida with other Mesozoic tribosphenic mammals from the southern continents (Gondwana) as well as the monotremes, while others favor closer affinities with northern (Laurasian) tribosphenic mammals or specifically with placentals. At least five species are known from the Maastrichtian, including a yet undescribed species known from a nearly complete skeleton that may represent a completely new group of mammals. The gondwanathere Lavanify, known from two teeth, is most closely related to other gondwanatheres found in India and Argentina. Two other teeth may represent another gondwanathere or a different kind of mammal. One molar fragment is one of the few known remains of a multituberculate mammal from Gondwana and another has been interpreted as either a marsupial or a placental.

Holotheria infraclass of mammals

Holotheria are a diverse group of mammals that are descendants of the last common ancestor of Kuehneotherium and Theria.

Marsupionta is a hypothesised subclass within the Mammalia group. The existence of Marsupionta is a postulation by some researchers as a category devolving upon a notional unification between marsupials with the egg-laying monotremes. Under this suggested classification, placental mammals would be the sister subclass to Marsupionta. The Marsupionta hypothesis was proposed in 1947 by W.K. Gregory and has since been the subject of multiple studies. This merging of marsupials and monotremes into the hypothesized subclass of Marsupionta is contrary to the widespread belief that pouch and placental mammals share the common subclass Theria that excludes monotremes.

Zhelestidae is a lineage of extinct eutherian mammals. Occurring in the Late Cretaceous from the Turonian to the Maastrichtian, they were an extremely successful group, with representatives present in Europe, Asia, India, Africa and North America, ostensibly rendering them a cosmopolitan clade. They were specialised towards an herbivorous lifestyle and were in fact initially considered stem-ungulates, but the presence of epipubics and "archaic" dental characters render them as non-placental eutherians.

Zalambdalestidae family of mammals (fossil)

Zalambdalestidae is a clade of Asian eutherians occurring during the Late Cretaceous. Once classified as Glires, features like epipubic bones and various cranial elements have identified these animals as outside of Placentalia, representing thus a specialised clade of non-placental eutherians without any living descendants, and potentially rather different from modern placentals in at least reproductive anatomy.


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