Epitheria

Epitheria Temporal range: Late Cretaceous - Recent
European mole (Boreoeutheria)
African bush elephant (Afrotheria)
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Subclass:	Theriiformes
Infraclass:	Eutheria
Magnorder:	Epitheria
Orders and Clades
Superorder Afrotheria: Afrosoricida (Golden moles and tenrecs) Macroscelidea (Elephant shrews) Tubulidentata (Aardvark) Hyracoidea (Hyraxes) Proboscidea (Elephants) Sirenia (Manatees, Dugongs) Clade Boreoeutheria: Superorder Laurasiatheria: Chiroptera (Bats) Eulipotyphla (Shrews, Moles) Cetacea (Whales, dolphins) Artiodactyla (Ruminants, etc.) Perissodactyla (Horses, etc.) Pholidota (Scaly Anteaters) Carnivora (Carnivores) Superorder Euarchontoglires: Rodentia (Rodents) Lagomorpha (Rabbits, etc.) Dermoptera (Flying Lemurs) Scandentia (Tree Shrews) Primates

Epitherians comprise all the placental mammals except the Xenarthra. They are primarily characterized by having a stirrup-shaped stapes in the middle ear, which allows for passage of a blood vessel. This is in contrast to the column-shaped stapes found in marsupials, monotremes, and xenarthrans. They are also characterized by having a shorter fibula relative to the tibia.

Epitheria — like Xenarthra and Afrotheria — originated after the K-Pg boundary 66 million years ago, with the placental diversification occurring within the first hundred thousand years after the K-Pg event and the first modern placental orders began appearing 2–3 million years later.^[1] Epitheres are one of the most successful groups of animals.

The monophyly of Epitheria has been challenged by molecular phylogenetic studies.^[2] While preliminary analysis of a set of retroposons shared by both Afrotheria, and Boreoeutheria (presence/absence data) supported the Epitheria clade,^[3] more extensive analysis of such transposable element insertions around the time of the divergence of Xenarthra, Afrotheria, and Boreoeutheria strongly support the hypothesis of a near-concomitant origin (trifurcation) of these three superorders of mammals.^[4][5]

Another analysis suggests that the root of this clade lies between the Atlantogenata and Boreoeutheria.^[6]

Placentalia	Xenarthra   Epitheria   Afrotheria   Boreoeutheria   Euarchontoglires Laurasiatheria

Alternative hypotheses

Alternative hypotheses place either Atlantogenata and Boreoeutheria, or Afrotheria and Exafroplacentalia (Notolegia) at the base of the tree:

Placentalia	Boreoeutheria   Euarchontoglires Laurasiatheria   Atlantogenata   Xenarthra Afrotheria

Placentalia	Afrotheria   Exafroplacentalia   Xenarthra   Boreoeutheria   Euarchontoglires Laurasiatheria

One Bayesian analysis places the root between Atlantogenata and Boreoeutheria.^[7]

References

1. O'Leary, M. A.; Bloch, J. I.; Flynn, J. J.; Gaudin, T. J.; Giallombardo, A.; Giannini, N. P.; Goldberg, S. L.; Kraatz, B. P.; Luo, Z.-X.; Meng, J.; Ni, X.; Novacek, M. J.; Perini, F. A.; Randall, Z. S.; Rougier, G. W.; Sargis, E. J.; Silcox, M. T.; Simmons, N. B.; Spaulding, M.; Velazco, P. M.; Weksler, M.; Wible, J. R.; Cirranello, A. L. (2013). "The Placental Mammal Ancestor and the Post-K-Pg Radiation of Placentals". Science. 339 (6120): 662–667. doi:10.1126/science.1229237. ISSN 0036-8075.
2. For example: Springer, Mark S., Michael J. Stanhope, Ole Madsen, and Wilfried W. de Jong. 2004. Molecules consolidate the placental mammal tree" Trends in Ecology and Evolution 19:430–438.
3. Kriegs, Jan Ole, Gennady Churakov, Martin Kiefmann, Ursula Jordan, Juergen Brosius, and Juergen Schmitz. (2006) Retroposed Elements as archives for the evolutionary history of placental mammals" PLoS Biol 4(4) e91.
4. Nishihara, H., Maruyama, S. & Okada, N. 2009. Retroposon analysis and recent geological data suggest near-simultaneous divergence of these three superorders of mammals. PNAS 106: 5235-40.
5. Churakov, G., Kriegs, J.O., Baertsch, R., Zemann, A., Brosius, J. & Schmitz, J. 2009. Mosaic retroposon insertion patterns in placental mammals. Genome Research 19: 868-75.
6. Song S, Liu L, Edwards SV, Wu S (2012) Resolving conflict in eutherian mammal phylogeny using phylogenomics and the multispecies coalescent model. Proc Natl Acad Sci USA
7. Morgan CC, Foster PG, Webb AE, Pisani D, McInerney JO, O'Connell MJ (2013) Heterogeneous models place the root of the placental mammal phylogeny" Mol Biol Evol 30(9) 2145-256 doi:10.1093/molbev/mst117

External links

• Summary of molecular support for Epitheria
• Waddell PJ, Kishino H, Ota R. 2001. A phylogenetic foundation for comparative mammalian genomics. Genome Inform Ser Workshop Genome Inform 12: 141–154
• Mark S. Springer, William J. Murphy, Eduardo Eizirik, and Stephen J. O'Brien (Edited by Morris Goodman). 2002 Placental mammal diversification and the Cretaceous-Tertiary boundary
• Wildman D.E.; Chen C.; Erez O.; Grossman L.I.; Goodman M.; Romero R. 2006. Evolution of the mammalian placenta revealed by phylogenetic analysis" PNAS 103 (9) 3203–3208
• Nikolaev, S., Montoya-Burgos, J.I., Margulies, E.H., Rougemont, J., Nyffeler, B., Antonarakis, S.E. 2007. Early history of mammals is elucidated with the ENCODE multiple species sequencing data" PLoS Genet 3:e2, doi:10.1371/journal.pgen.0030002.
• Gennady Churakov, Jan Ole Kriegs, Robert Baertsch, Anja Zemann, Jürgen Brosius, Jürgen Schmitz. 2008. Mosaic retroposon insertion patterns in placental mammals