Eutheria (from Greek εὐ-, eú- 'good, right' and θηρίον, thēríon 'beast'; lit.'true beasts'), also called Pan-Placentalia, is the clade consisting of placentals and all therian mammals that are more closely related to placentals than to marsupials.
Eutherians are distinguished from non-eutherians 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,[1] though epipubic bones are present in many primitive eutherians.[2] Eutheria was named in 1872 by Theodore Gill; in 1880, Thomas Henry Huxley defined it to encompass a more broadly defined group than Placentalia.[3]
The earliest unambiguous eutherians are known from the Early Cretaceous Yixian Formation of China, dating around 120 million years ago.[4] Two tribosphenic mammals, Durlstodon and Durlstotherium from the Berriasian age (~145-140 million years ago) of the Early Cretaceous in southern England have also been suggested to represent early eutherians.[5][6] Another possible eutherian species Juramaia sinensis has been dated at 161 million years ago from the early Late Jurassic (Oxfordian) of China.[7] However some authors have considered Juramaia as a stem therian instead,[5][6] and some sources have doubted the dating of the specimen.[8]
Characteristics
Distinguishing features are:
an enlarged malleolus ("little hammer") at the bottom of the tibia, the larger of the two shin bones[9]
the joint between the first metatarsal bone and the entocuneiform bone (the innermost of the three cuneiform bones) in the foot is offset farther back than the joint between the second metatarsal and middle cuneiform bones—in metatherians these joints are level with each other[9]
various features of jaws and teeth[9] including: having three molars in the halves of each jaw, each upper canine having two roots, the paraconid on the last lower premolar is pronounced, the talonid region of the lower molars is narrower than the trigonid.[10]
Some older systems contained an order called Cimolesta (sensu lato), which contains the above taxa Cimolestidae, Taeniodonta and Didymoconidae, but also (all or some of) the taxa †Ptolemaiidae, †Palaeoryctidae, †Wyolestidae, †Pantolesta (probably inclusive of the family †Horolodectidae), †Tillodontia, †Apatotheria, †Pantodonta, Pholidota and †Palaeanodonta. Those additional taxa (all of which are usually considered members of Placentalia sensu stricto today) were thus also placed among basal Eutheria in such older systems and were placed next to Cimolestidae.
Some systems also included the †Creodonta and/or †Dinocerata as basal Eutherians.
Some authors classify the taxa, which are at the end of the above system of basal Eutheria, as part of Placentalia sensu stricto. More specifically, depending on the author, this applies to the taxa of the above system that are placed from (and inclusive of) Leptictida or Asioryctitheria or Adapisoriculidae down to (and inclusive of) Oxyprimus.
Evolutionary history
Eutheria contains several extinctgenera 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 outdated placental group Insectivora, while zhelestids have been considered primitive ungulates.[33] However, more recent studies have suggested these enigmatic taxa represent stem group eutherians, more basal to Placentalia.[34][35]
The weakly favoured cladogram favours Boreoeutheria as a basal eutherian clade as sister to the Atlantogenata.[36][37][38]
Many non-placental eutherians are thought to have been insectivores, as is the case with many primitive mammals.[41] However, the zhelestids are thought to have been herbivorous.[40] Body size of eutherians was generally small during the Cretaceous period, but the range of body sizes increased dramatically after the K-Pg extinction, predominantly among placentals.[42]
^Rook, Deborah L.; Hunter, John P. (2013). "Rooting Around the Eutherian Family Tree: the Origin and Relations of the Taeniodonta". Journal of Mammalian Evolution. 21: 1–17. doi:10.1007/s10914-013-9230-9.
^Wilson Mantilla, Gregory P.; Renne, Paul R.; Samant, Bandana; Mohabey, Dhananjay M.; Dhobale, Anup; Tholt, Andrew J.; Tobin, Thomas S.; Widdowson, Mike; Anantharaman, S.; Dassarma, Dilip Chandra; Wilson Mantilla, Jeffrey A. (2022). "New mammals from the Naskal intertrappean site and the age of India's earliest eutherians". Palaeogeography, Palaeoclimatology, Palaeoecology. 591. Elsevier BV. Bibcode:2022PPP...59110857W. doi:10.1016/j.palaeo.2022.110857. ISSN0031-0182.
^Scott, Craig S (2019-01-18). "Horolodectidae: a new family of unusual eutherians (Mammalia: Theria) from the Palaeocene of Alberta, Canada". Zoological Journal of the Linnean Society. 185 (2): 431–458. doi:10.1093/zoolinnean/zly040. ISSN0024-4082.
^Kynigopoulou, Zoi; Brusatte, Stephen; Fraser, Nicholas; Wood, Rachel; Williamson, Tom; Shelley, Steve (2023-06-12). Phylogeny, evolution, and anatomy of Taeniodonta (Mammalia: Eutheria) and implications for the mammalian evolution after the Cretaceous-Palaeogene mass extinction (Thesis). University Of Edinburgh. doi:10.7488/ERA/3414. hdl:1842/40653.
^Rose, Kenneth D. (2006). The beginning of the age of mammals. Baltimore: Johns Hopkins University Press. ISBN9780801892219.
^Wible, J. R.; Rougier, G. W.; Novacek, M. J.; Asher, R. J. (2007). "Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary". Nature. 447 (7147): 1003–1006. Bibcode:2007Natur.447.1003W. doi:10.1038/nature05854. PMID17581585.
^ abGheerbrant, Emmanuel; Teodori, Dominique (2021-03-24). "An enigmatic specialized new eutherian mammal from the Late Cretaceous of Western Europe (Northern Pyrenees)". Comptes Rendus Palevol (13). doi:10.5852/cr-palevol2021v20a13. ISSN1777-571X.