Metatheria

Metatheria Temporal range:
Lycopsis longirostris, an extinct sparassodont, a relative of the marsupials
A mouse opossum (Marmosa)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Subclass:	Theria
Clade:	Metatheria Huxley, 1880
Subgroups
†?Sinodelphys †Holoclemensia †Adinodon †Deltatheroida Marsupialiformes †Adelodelphys †Alphadontidae †Asiadelphia †Sinbadelphys †Camptomus? †Archimetatheria †Stagodontidae †Iqualadelphis †Sparassodonta †Pucadelphyidae †Herpetotheriidae †Anatoliadelphyidae †Polydolopimorphia †Peradectidae †Pediomyidae Marsupialia

Metatheria (from Greek μετά-, metá- 'changed' and θηρίον, thēríon 'beast'; lit. 'changed beasts') 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 more inclusive group than the marsupials; it contains all marsupials as well as many extinct non-marsupial relatives. It is one of two groups placed in the clade Theria alongside Eutheria, which contains the placentals. Remains of metatherians have been found on all of Earths continents.

Description

Distinctive characteristics (synapomorphies) of Metatheria include: a prehensile tail, the development of a capitular tail on the humerus, the loss of tooth replacement on the second and fifth premolars and the retention of decidious teeth on the lower fifth premolars, the lower canines outwardly diverge from each other, the angular process on the dentary is equal to or less than half the length of the ramus, the dentary has a posterior masseteric shelf, and the lower 5th premolar has a "very trenchant" cristid obliqua/ectolophid. The permanent deciduous lower fifth premolars are molar like and were historically identified as 1st molars, with the third premolar found in basal therians being lost, leaving 4 premolars in the halves of each jaw.

Evolutionary history

The relationships between the three extant divisions of mammals (monotremes, marsupials, and placentals) was long a matter of debate among taxonomists. Most morphological evidence comparing traits, such as the number and arrangement of teeth and the structure of the reproductive and waste elimination systems, favors a closer evolutionary relationship between marsupials and placentals than either has with the monotremes, as does most genetic and molecular evidence.

The earliest possible known metatherian is Sinodelphys szalayi, which lived in China during the Early Cretaceous around 125 million years ago (mya). This makes it a contemporary to some early eutherian species that have been found in the same area. However, Bi et al. (2018) reinterpreted Sinodelphys as an early member of Eutheria. The oldest uncontested metatherians are now 110 million year old fossils from western North America. Metatherians were widespread in Asia and North America during the Late Cretaceous, including both Deltatheroida and Marsupialiformes, with fossils also known from Europe during this time. During the Late Cretaceous, metatherians were more diverse than eutherians in North America. Metatherians underwent a severe decline during the Cretaceous–Paleogene extinction event, more severe than that suffered by contemporary eutherians and multituberculates, and were slower to recover diversity.

Morphological and species diversity of metatherians in Laurasia remained low in comparison to eutherians throughout the Cenozoic. The two major groups of Cenozoic Laurasian metatherians, the opossum-like herpetotheriids and peradectids persisted into the Miocene before becoming extinct, with the North American herpetotheriid Herpetotherium, the European herpetotheriid Amphiperatherium and the peradectids Siamoperadectes and Sinoperadectes from Asia being the youngest Laurasian non-marsupial metatherians (with marsupials invading North America during the Pliocene-Pleistocene as part of the Great American interchange). Metatherians first arrived in Afro-Arabia during the Paleogene, probably from Europe, including the possible peradectoid Kasserinotherium from the Early Eocene of Tunisia and the herpetotheriid Peratherium africanum from the Early Oligocene of Egypt and Oman. The youngest African metatherian is the possible herpetotheriid Morotodon from the late Early Miocene of Uganda.

Metatherians arrived in South America from North America during the latest Cretaceous or Paleocene and underwent a major diversificiation, with South American metatherians including both the ancestors of extant marsupials as well as the extinct Sparassodonta, which were major predators in South American ecosystems during most of the Cenozoic, up until their extinction in the Pliocene, as well as the Polydolopimorphia, which likely had a wide range of diets. Metatherians then declined in diversity in South America during the Late Eocene as well as the later Oligocene epoch. The oldest known Australian marsupials are from the early Eocene, and are thought to have arrived in the region after having dispersed via Antarctica from South America. During the Oligocene epoch, Australian metatherians radiated rapidly, which contributed most to the global peak in metatherian diversity during the Early Miocene. The only known Antarctic metatherians are from the Early Eocene La Meseta Formation of the Antarctic Peninsula, where they are the most diverse group of mammals, and include marsupials as well as polydolopimorphians.

Classification

Below is a metatherian cladogram from Wilson et al. (2016):

Metatheria

Holoclemensia Deltatheroida Pappotheriidae Pappotherium Deltatheridiidae Sulestes Oklatheridium Tsagandelta Lotheridium Deltatheroides Deltatheridium Nanocuris Atokatheridium Marsupialiformes Gurlin Tsav skull Borhyaenidae Mayulestes Jaskhadelphys Andinodelphys Pucadelphys Asiatherium Iugomortiferum Kokopellia Aenigmadelphys Anchistodelphys Glasbiidae Glasbius Pediomyidae Pediomys Stagodontidae Pariadens Eodelphis Didelphodon Turgidodon Alphadon Alphadontidae Albertatherium Marsupialia

Cladogram after:

Metatheria	Deltatheriidae Kokopellia Asiatherium Peradectidae Stagodontidae Pucadelphyidae Sparassodonta Amphiperatherium Peratherium "Herpetotheriidae" Herpetotherium Marsupialia

Below is a listing of metatherians that do not fall readily into well-defined groups.

Basal Metatheria

• †Archaeonothos henkgodthelpi Beck 2015
• †Esteslestes ensis Novacek et al. 1991
• †Ghamidtherium dimaiensis Sánches-Villagra et al. 2007
• †Kasserinotherium tunisiense Crochet 1989
• †Palangania brandmayri Goin et al. 1998
• †Perrodelphys coquinense Goin et al. 1999

Ameridelphia incertae sedis:

• †Apistodon exiguus (Fox 1971) Davis 2007
• †Cocatherium lefipanum Goin et al. 2006
• †Dakotadens morrowi Eaton 1993
• †Iugomortiferum thoringtoni Cifelli 1990b
• †Marambiotherium glacialis Goin et al. 1999
• †Marmosopsis juradoi Paula Couto 1962 [Marmosopsini Kirsch & Palma 1995]
• †Pascualdelphys fierroensis
• †Progarzonia notostylopense Ameghino 1904
• †Protalphadon Cifelli 1990
  • †P. lulli (Clemens 1966) Cifelli 1990a
  • †P. foxi Johnson 1996

Marsupialia incertae sedis:

• †Itaboraidelphys camposi Marshall & de Muizon 1984
• †Mizquedelphys pilpinensis Marshall & de Muizon 1988
• †Numbigilga ernielundeliusi Beck et al. 2008 {Numbigilgidae Beck et al. 2008}