Paenungulatomorpha

Paenungulatomorpha; Temporal range: Paleocene - Recent, 61–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N
	Crowned paenungulates and Ocepeia
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Superorder:	Afrotheria
Clade:	Paenungulatomorpha; Gheerbrant, 2016
Subgroups
	†Hadrogeneios; †Ocepeia; †Abdounodus; Paenungulata;

Paenungulatomorpha is a clade of afrotherian mammals that can be characterized according to Gheerbrant et al. (2016)^[1]:

by a mandibular retromolar fossa, the absence of hypocone, an ectoloph selenodont and linked to strong styles such as mesostyle in basal taxa, and a more or less developed pseudohypocone.

Originally it was thought paenungulates were closely related to the perissodactyls in the clade Altungulata due to shared attributes in the dental, osteological and soft-tissues.^[2]^[3]^[4] However this view was contested as molecular studies have widely supported that paenungulates are not related to the perissodactyls, instead more related to the widely different clade Afroinsectiphilia, with them forming the clade Afrotheria.^[5]^[6]^[7]^[8]^[9] There has been also some morphological data to support Afrotheria,^[10]^[11]^[12] though the paleontological record to support was lacking until reexamination of the genera Ocepeia^[13] and Abdounodus,^[1] as well as the description of the basal Hadrogeneios in 2023^[14] has shown not only fossil evidence for the inclusion of paenungulates as afrotherians but also any morphological similarities between paenungulates and perissodactyls is due to convergence.^[1]

Below is a cladogram from Gheerbrant (2023):^[14]

Paenungulatomorpha

†Hadrogeneios

†Ocepeia

†Abdounodus

	Sirenia

	Proboscidea

References

^ ^a ^b ^c Emmanuel Gheerbrant; Andrea Filippo; Arnaud Schmitt (2016). "Convergence of Afrotherian and Laurasiatherian Ungulate-Like Mammals: First Morphological Evidence from the Paleocene of Morocco". PLOS ONE. 11 (7): e0157556. Bibcode:2016PLoSO..1157556G. doi:10.1371/journal.pone.0157556. PMC 4934866. PMID 27384169.
^ Fischer, Martin S. (1986). "Die Stellung der Schliefer (Hyracoidea) im phylogenetischen System der Eutheria". Courier Forschungsinstitut Senckenberg. 84: 1–132. ISBN 9783924500214. OCLC 15682192.
^ Prothero, Donald R.; Schoch, Robert M. (1989). "Origin and Evolution of the Perissodactyla: Summary and Synthesis" (PDF). In Prothero, Donald R.; Schoch, Robert M. (eds.). The Evolution of Perissodactyls. New York: Oxford University Press. pp. 504–37. ISBN 9780195060393. OCLC 19268080. Retrieved 5 May 2013.
^ Rose, Kenneth David (2006). The beginning of the age of mammals. Baltimore: JHU Press. ISBN 0801884721.
^ Stanhope, M. J.; Waddell, V. G.; Madsen, O.; de Jong, W.; Hedges, S. B.; Cleven, G. C.; Kao, D.; Springer, M. S. (1998). "Molecular evidence for multiple origins of Insectivora and for a new order of endemic African insectivore mammals". Proceedings of the National Academy of Sciences. 95 (17): 9967–9972. Bibcode:1998PNAS...95.9967S. doi:10.1073/pnas.95.17.9967. PMC 21445. PMID 9707584.
^ Springer, Mark S.; Michael J. Stanhope; Ole Madsen; Wilfried W. de Jong (2004). "Molecules consolidate the placental mammal tree" (PDF). Trends in Ecology & Evolution. 19 (8): 430–438. doi:10.1016/j.tree.2004.05.006. PMID 16701301.
^ Robinson, T. J.; Fu, B.; Ferguson-Smith, M. A.; Yang, F. (2004). "Cross-species chromosome painting in the golden mole and elephant-shrew: support for the mammalian clades Afrotheria and Afroinsectiphillia but not Afroinsectivora". Proceedings of the Royal Society B: Biological Sciences. 271 (1547): 1477–1484. doi:10.1098/rspb.2004.2754. PMC 1691750. PMID 15306319.
^ Nishihara, H.; Satta, Y.; Nikaido, M.; Thewissen, J.G.M.; Stanhope, M.J.; Okada, N. (2005). "A retroposon analysis of Afrotherian phylogeny". Molecular Biology and Evolution. 22 (9): 1823–1833. doi:10.1093/molbev/msi179. PMID 15930154.
^ Scornavacca C, Belkhir K, Lopez J, Dernat R, Delsuc F, Douzery EJ, Ranwez V (April 2019). "OrthoMaM v10: Scaling-up orthologous coding sequence and exon alignments with more than one hundred mammalian genomes". Molecular Biology and Evolution. 36 (4): 861–862. doi:10.1093/molbev/msz015. PMC 6445298. PMID 30698751.
^ Tabuce, R.; Marivaux, L.; Adaci, M.; Bensalah, M.; Hartenberger, J.-L.; Mahboubi, M.; Mebrouk, F.; Tafforeau, P.; Jaeger, J.-J. (2007). "Early Tertiary mammals from North Africa reinforce the molecular Afrotheria clade". Proceedings of the Royal Society B: Biological Sciences. 274 (1614): 1159–1166. doi:10.1098/rspb.2006.0229. PMC 2189562. PMID 17329227.
^ Seiffert, Erik R (2007). "A new estimate of afrotherian phylogeny based on simultaneous analysis of genomic, morphological, and fossil evidence". BMC Evolutionary Biology. 7 (1): 224. Bibcode:2007BMCEE...7..224S. doi:10.1186/1471-2148-7-224. PMC 2248600. PMID 17999766.
^ Sánchez-Villagra, Marcelo R.; Narita, Yuichi; Kuratani, Shigeru (2007). "Thoracolumbar vertebral number: The first skeletal synapomorphy for afrotherian mammals". Systematics and Biodiversity. 5 (1): 1–7. Bibcode:2007SyBio...5Q...1S. doi:10.1017/S1477200006002258. S2CID 85675984.
^ Emmanuel Gheerbrant; Mbarek Amaghzaz; Baadi Bouya; Florent Goussard; Charlène Letenneur (2014). "Ocepeia (Middle Paleocene of Morocco): The Oldest Skull of an Afrotherian Mammal". PLOS ONE. 9 (2): e89739. Bibcode:2014PLoSO...989739G. doi:10.1371/journal.pone.0089739. PMC 3935939. PMID 24587000.
^ ^a ^b Gheerbrant, Emmanuel (2023-04-27). "Ancestral radiation of paenungulate mammals (Paenungulatomorpha)—new evidence from the Paleocene of Morocco". Journal of Vertebrate Paleontology. 42 (5). doi:10.1080/02724634.2023.2197971. ISSN 0272-4634.

[Gheerbrantetal2016-1] Emmanuel Gheerbrant; Andrea Filippo; Arnaud Schmitt (2016). "Convergence of Afrotherian and Laurasiatherian Ungulate-Like Mammals: First Morphological Evidence from the Paleocene of Morocco". PLOS ONE. 11 (7): e0157556. Bibcode:2016PLoSO..1157556G. doi:10.1371/journal.pone.0157556. PMC 4934866. PMID 27384169.

[Fischer1986-2] Fischer, Martin S. (1986). "Die Stellung der Schliefer (Hyracoidea) im phylogenetischen System der Eutheria". Courier Forschungsinstitut Senckenberg. 84: 1–132. ISBN 9783924500214. OCLC 15682192.

[Prothero&Schoch1989-3] Prothero, Donald R.; Schoch, Robert M. (1989). "Origin and Evolution of the Perissodactyla: Summary and Synthesis" (PDF). In Prothero, Donald R.; Schoch, Robert M. (eds.). The Evolution of Perissodactyls. New York: Oxford University Press. pp. 504–37. ISBN 9780195060393. OCLC 19268080. Retrieved 5 May 2013.

[Rose2006-4] Rose, Kenneth David (2006). The beginning of the age of mammals. Baltimore: JHU Press. ISBN 0801884721.

[Stanhope-5] Stanhope, M. J.; Waddell, V. G.; Madsen, O.; de Jong, W.; Hedges, S. B.; Cleven, G. C.; Kao, D.; Springer, M. S. (1998). "Molecular evidence for multiple origins of Insectivora and for a new order of endemic African insectivore mammals". Proceedings of the National Academy of Sciences. 95 (17): 9967–9972. Bibcode:1998PNAS...95.9967S. doi:10.1073/pnas.95.17.9967. PMC 21445. PMID 9707584.

[Springeretal2004-6] Springer, Mark S.; Michael J. Stanhope; Ole Madsen; Wilfried W. de Jong (2004). "Molecules consolidate the placental mammal tree" (PDF). Trends in Ecology & Evolution. 19 (8): 430–438. doi:10.1016/j.tree.2004.05.006. PMID 16701301.

[Robinsonetal2004-7] Robinson, T. J.; Fu, B.; Ferguson-Smith, M. A.; Yang, F. (2004). "Cross-species chromosome painting in the golden mole and elephant-shrew: support for the mammalian clades Afrotheria and Afroinsectiphillia but not Afroinsectivora". Proceedings of the Royal Society B: Biological Sciences. 271 (1547): 1477–1484. doi:10.1098/rspb.2004.2754. PMC 1691750. PMID 15306319.

[8] Nishihara, H.; Satta, Y.; Nikaido, M.; Thewissen, J.G.M.; Stanhope, M.J.; Okada, N. (2005). "A retroposon analysis of Afrotherian phylogeny". Molecular Biology and Evolution. 22 (9): 1823–1833. doi:10.1093/molbev/msi179. PMID 15930154.

[Scornavacca2019-9] Scornavacca C, Belkhir K, Lopez J, Dernat R, Delsuc F, Douzery EJ, Ranwez V (April 2019). "OrthoMaM v10: Scaling-up orthologous coding sequence and exon alignments with more than one hundred mammalian genomes". Molecular Biology and Evolution. 36 (4): 861–862. doi:10.1093/molbev/msz015. PMC 6445298. PMID 30698751.

[Tabuce2007-10] Tabuce, R.; Marivaux, L.; Adaci, M.; Bensalah, M.; Hartenberger, J.-L.; Mahboubi, M.; Mebrouk, F.; Tafforeau, P.; Jaeger, J.-J. (2007). "Early Tertiary mammals from North Africa reinforce the molecular Afrotheria clade". Proceedings of the Royal Society B: Biological Sciences. 274 (1614): 1159–1166. doi:10.1098/rspb.2006.0229. PMC 2189562. PMID 17329227.

[Seifert2007-11] Seiffert, Erik R (2007). "A new estimate of afrotherian phylogeny based on simultaneous analysis of genomic, morphological, and fossil evidence". BMC Evolutionary Biology. 7 (1): 224. Bibcode:2007BMCEE...7..224S. doi:10.1186/1471-2148-7-224. PMC 2248600. PMID 17999766.

[SanchezVillagra-12] Sánchez-Villagra, Marcelo R.; Narita, Yuichi; Kuratani, Shigeru (2007). "Thoracolumbar vertebral number: The first skeletal synapomorphy for afrotherian mammals". Systematics and Biodiversity. 5 (1): 1–7. Bibcode:2007SyBio...5Q...1S. doi:10.1017/S1477200006002258. S2CID 85675984.

[Ocepeia-13] Emmanuel Gheerbrant; Mbarek Amaghzaz; Baadi Bouya; Florent Goussard; Charlène Letenneur (2014). "Ocepeia (Middle Paleocene of Morocco): The Oldest Skull of an Afrotherian Mammal". PLOS ONE. 9 (2): e89739. Bibcode:2014PLoSO...989739G. doi:10.1371/journal.pone.0089739. PMC 3935939. PMID 24587000.

[Gheerbrant2023-14] Gheerbrant, Emmanuel (2023-04-27). "Ancestral radiation of paenungulate mammals (Paenungulatomorpha)—new evidence from the Paleocene of Morocco". Journal of Vertebrate Paleontology. 42 (5). doi:10.1080/02724634.2023.2197971. ISSN 0272-4634.

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