Pholidotamorpha

Pholidotamorphs Temporal range: Paleocene–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Various Pholidotamorph genera; clockwise from top left: Manis, Ernanodon, Xenocranium, Metacheiromys, Eurotamandua, Eomanis.
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Mirorder:	Ferae
Clade:	Pholidotamorpha Gaudin et al., 2009[1]
Orders
Pholidota †Palaeanodonta

Pholidotamorpha ("pangolin-like forms") is a clade of placental, mostly ant- and termite-eating mammals that (partially) physically resemble the anteaters or armadillos. However, those aforementioned species are now placed in the order Xenarthra, along with sloths; Pholidotamorpha is now classified under the mirorder Ferae, which includes the order Carnivora (carnivorous mammals) and the pangolins (Pholidota) as well as the prehistoric order Palaeanodonta, containing only extinct species.[1]

Classification and phylogeny

History of taxonomy

Both the Pholidota and Palaeanodonta orders were formerly placed with other orders of ant-eating mammals, most notably Xenarthra (armadillos, sloths, anteaters, which they superficially resemble); some palaeontologists, throughout the history of zoology, have placed pangolins and palaeanodonts as a suborder, Pholidota, in the greater order Cimolesta, alongside the extinct family Ernanodontidae as a separate suborder Ernanodonta near it. However, this idea fell out-of-favor when it was determined that cimolestids were not truly placental mammals.[2]

Newer genetic evidence indicates instead that the closest living relatives to Pholidota are the members of order Carnivora, together forming the mirorder Ferae.[3][4][5] In 2009, pangolins and palaeanodonts were together placed within the clade Pholidotamorpha.[1] A 2012 study of new remains, found in Late Paleocene Mongolian strata, have led to the assessment that extinct genus Ernanodon is closely-related to another extinct genus, Metacheiromys, and is a member of the extinct order Palaeanodonta.[6]

Classification

Clade: Pholidotamorpha (Gaudin, 2009) Order: Pholidota (Weber, 1904) (pangolins) Suborder: Eupholidota (Gaudin, 2009) (true pangolins) Family: †Eurotamanduidae (Szalay & Schrenk, 1994) Genus: †Euromanis (Gaudin, 2009) Incertae sedis †Pholidota sp. [BC 16’08] (Pickford, 2008) Order: †Palaeanodonta (Matthew, 1918) (stem-pangolins) Family: †Epoicotheriidae ^(paraphyletic family) (Simpson, 1927) Family: †Ernanodontidae (Ding, 1979) Family: †Escavadodontidae (Rose & Lucas, 2000) Family: †Metacheiromyidae ^(paraphyletic family) (Wortman, 1903) Incertae sedis: Genus: †Arcticanodon (Rose, 2004) Genus: †Melaniella (Fox, 1984)

Phylogeny

The phylogenetic relationships of clade Pholidotamorpha are shown in the following cladogram:[7][8][1][6][9][10][11]

Ferae	Pan-Carnivora  Pholidotamorpha   Pholidota  Eupholidota †Eurotamanduidae  ?  †Pholidota sp. (BC 16’08) †Euromanis  sensu stricto   †Palaeanodonta  †Escavadodontidae  ?  †Melaniella †Amelotabes †Epoicotheriidae †Epoicotherium †Molaetherium †Xenocranium †Epoicotheriinae †Tetrapassalus †Dipassalus †Alocodontulum †Auroratherium †Pentapassalus †Tubulodon  ?  †Arcticanodon †Propalaeanodontinae †Mylanodon †Brachianodon †Palaeanodon †Metacheiromyinae  †Metacheiromys †Ernanodontidae  (Pholidota sensu lato)	†Epoicotherium/Xenocranium clade †Metacheiromyidae

See also

• Mammal classification
• Ferae

References

1. Gaudin, Timothy (2009). "The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis" (PDF). Journal of Mammalian Evolution. 16 (4). Heidelberg, Germany: Springer Science+Business Media: 235–305. doi:10.1007/s10914-009-9119-9. S2CID 1773698. Archived from the original (PDF) on 2015-09-25. Retrieved 2020-08-05.
2. Rook, D. L.; Hunter, J. 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. S2CID 17074668.
3. Murphy, Willian J., et al. (2001-12-14). "Resolution of the Early Placental Mammal Radiation Using Bayesian Phylogenetics". Science. 294 (5550): 2348–2351. Bibcode:2001Sci...294.2348M. doi:10.1126/science.1067179. PMID 11743200. S2CID 34367609.
4. Beck, Robin MD; Bininda-Emonds, Olaf RP; Cardillo, Marcel; Liu, Fu-Guo; Purvis, Andy (2006). "A higher-level MRP supertree of placental mammals". BMC Evolutionary Biology. 6 (1): 93. doi:10.1186/1471-2148-6-93. PMC 1654192. PMID 17101039.
5. Mark S Springer, Christopher A Emerling, John Gatesy, Jason Randall, Matthew A. Collin, Nikolai Hecker, Michael Hiller, Frédéric Delsuc (2019) Odontogenic ameloblast-associated (ODAM) is inactivated in toothless/enamelless placental mammals and toothed whales
6. Kondrashov, Peter; Agadjanian, Alexandre K. (2012). "A nearly complete skeleton of Ernanodon (Mammalia, Palaeanodonta) from Mongolia: morphofunctional analysis". Journal of Vertebrate Paleontology. 32 (5): 983–1001. doi:10.1080/02724634.2012.694319. ISSN 0272-4634. S2CID 86059673.
7. Amrine-madsen, H.; Koepfli, K.P.; Wayne, R.K.; Springer, M.S. (2003). "A new phylogenetic marker, apolipoprotein B, provides compelling evidence for eutherian relationships". Molecular Phylogenetics and Evolution. 28 (2): 225–240. doi:10.1016/S1055-7903(03)00118-0. PMID 12878460.
8. Kenneth D. Rose (2008). "Palaeanodonta and Pholidota". 9 - Palaeanodonta and Pholidota. pp. 135–146. doi:10.1017/CBO9780511541438.010. ISBN 9780521781176.
9. Halliday, Thomas J. D.; Upchurch, Paul; Goswami, Anjali (2015). "Resolving the relationships of Paleocene placental mammals" (PDF). Biological Reviews. 92 (1): 521–550. doi:10.1111/brv.12242. ISSN 1464-7931. PMC 6849585. PMID 28075073.
10. Solé, Floréal; Ladevèze, Sandrine (2017). "Evolution of the hypercarnivorous dentition in mammals (Metatheria,Eutheria) and its bearing on the development of tribosphenic molars". Evolution & Development. 19 (2): 56–68. doi:10.1111/ede.12219. PMID 28181377. S2CID 46774007.
11. Prevosti, F. J., & Forasiepi, A. M. (2018). "Introduction. Evolution of South American Mammalian Predators During the Cenozoic: Paleobiogeographic and Paleoenvironmental Contingencies"