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Temporal range: 82.0–0 Ma Late Cretaceous - present[1]
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Scrotifera
Grandorder: Ferungulata
Simpson, 1945[2]
  • Cetferungulata (Arnason, 1999)[3]
  • Fereuungulata (Waddell, 1999)[4]

Ferungulata ("wild beasts and ungulates") is a grandorder of placental mammals that groups together mirorder Ferae and clade Pan-Euungulata. It has existed in two guises, a traditional one based on morphological analysis and a revised one taking into account more recent molecular analyses. The Fereungulata is a sister group to the order Chiroptera (bats) and together they make clade Scrotifera.

The traditional Ferungulata was established by George Gaylord Simpson in 1945.[2] It grouped together the extant orders Carnivora, Perissodactyla and Artiodactyla with the Tubulidentata and the superorder Paenungulata, as well as a number of orders known only from fossils.

Simpson established the grouping on the basis of morphological criteria, but this traditional understanding of Ferungulata has been challenged by a more recent classification, relying upon genetic criteria.[4][1][5] These studies separated his ungulate orders into two distinct placental groups, within Afrotheria and Laurasiatheria, respectively. The 'true' ungulates (mirorder Euungulata), Perissodactyla and Artiodactyla, are included in the revised group, along with the Carnivora, and with the addition of pangolins (order Pholidota), but the Tubulidentata and paenungulates are excluded. Although Simpson placed whales (Cetacea) in a separate cohort, recent evidence linking them to Artiodactyla would mean that they belong here as well. To reflect this difference, the revised clade is usually referred to as Fereungulata.

Phylogeny within grandorder Ferungulata[6][7][8][9][5]

Euarchontoglires Bruno Liljefors - Hare studies 1885 white background.jpg


Eulipotyphla Erinaceus europaeus - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg


Chiroptera Braunes Langohr (Plectus auritus).jpg


Euungulata Equus quagga (white background).jpg




Pholidotamorpha Pangolin Hardwicke (white background).jpg

Pan-Carnivora Dogs, jackals, wolves, and foxes (Plate XI).jpg

The cladogram has been reconstructed from mitochondrial and nuclear DNA and protein characters, as well as the fossil record.

See also[edit]


  1. ^ a b Zhou, X.; Xu, S.; Xu, J.; Chen, B.; Zhou, K.; Yang, G. (2012). "Phylogenomic Analysis Resolves the Interordinal Relationships and Rapid Diversification of the Laurasiatherian Mammals". Systematic Biology. 61 (1): 150–164. doi:10.1093/sysbio/syr089. ISSN 1063-5157. PMC 3243735.
  2. ^ a b Simpson, George Gaylord (1945). "The principles of classification and a classification of mammals". Bulletin of the American Museum of Natural History. 85: 350. hdl:2246/1104.
  3. ^ Arnason U., Gullberg A., Janke A. (1999.) "The mitochondrial DNA molecule of the aardvark, Orycteropus afer, and the position of the Tubulidentata in the eutherian tree." Proc R Soc Lond B 266:339–345
  4. ^ a b Waddell; et al. (1999). "Using Novel Phylogenetic Methods to Evaluate Mammalian mtDNA, Including Amino Acid-Invariant Sites-LogDet plus Site Stripping, to Detect Internal Conflicts in the Data, with Special Reference to the Positions of Hedgehog, Armadillo, and Elephant" (PDF). Systematic Biology. 48 (1): 31–53. doi:10.1080/106351599260427. PMID 12078643. Retrieved 4 October 2011.
  5. ^ a b Xue Lv, Jingyang Hu, Yiwen Hu, Yitian Li, Dongming Xu, Oliver A. Ryder, David M. Irwin, Li Yu (2021.) "Diverse phylogenomic datasets uncover a concordant scenario of laurasiatherian interordinal relationships", Molecular Phylogenetics and Evolution, Volume 157
  6. ^ O’Leary, M. A., Bloch JI, Flynn, J. J., Gaudin, T. J., Giallombardo, A., Giannini, N. P., Goldber, S. L, Kraatz, B. P., Luo, Z-X, Jin Meng, Xijun Ni, 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.
  7. ^ Burger, Benjamin J. (2015-10-15). The Systematic Position of the Saber-Toothed and Horned Giants of the Eocene: The Uintatheres (Order Dinocerata) (PDF). Society of Vertebrate Paleontology 75th Annual Meeting. Dallas. Retrieved 2020-02-20. Conference abstract (p. 99). Explanation and conclusions: Episode 17: Systematic position of the Uintatheres (Order Dinocerata) on YouTube.
  8. ^ Orliac, M. J.; o'Leary, M. A. (2016). "The inner ear of Protungulatum (Pan-Euungulata, Mammalia)". Journal of Mammalian Evolution. 23 (4): 337–352. doi:10.1007/s10914-016-9327-z. S2CID 33676627.
  9. ^ Frank Zachos (2020.) "Mammalian Phylogenetics: A Short Overview of Recent Advances"