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Condylarthra

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Condylarthra
Temporal range: Late Cretaceous to Late Eocene 66.3–37.2 Ma
Skull and vertebrae of Arctocyon primaevus
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Pan-Euungulata
Order: †Condylarthra
Cope, 1881
Families

Condylarthra, also known as archaic ungulates,[7][8] is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene epochs.[9] It is now largely considered to be a wastebasket taxon,[10] having served as a dumping ground for classifying ungulates which had not been clearly established as part of either Perissodactyla or Artiodactyla, being composed thus of several unrelated lineages.[11][12][10] Spharnotheriodontidae recovered as a family of Litopterna.[13] Additionally, some condylarths, such as Kharmerungulatum, were found to be have been outside of ungulates entirely.[14] Arctocyonidae was recovered as a polyphyletic family of mammals, with the placement within Ferungulata varying between genera.[15][16]

Taxonomy

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Condylarthra always was a problematic group. When first described by Cope 1881, Phenacodontidae was the type and only family therein. Cope 1885, however, raised Condylarthra to an order and included a wide range of diverse placentals with generalized dentitions and postcranial skeletons. More recent researchers (i.e. post-WW2) have been more restrictive; either including only a limited number of taxa, or proposing that the term should be abandoned altogether.[17] Due to their primitive characteristics condylarths have been considered ancestral to several ungulate orders, including the living Artiodactyla, Cetacea, Perissodactyla, Hyracoidea, Sirenia, and Proboscidea, as well as the extinct Desmostylia, Embrithopoda, Litopterna, Notoungulata, and Astrapotheria.[8]

Prothero, Manning & Fischer 1988 delimited condylarths as those having the following characters, but lacking the specializations present in more derived orders:[17]

Evolutionary history

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Ectocion, a small plant-eating phenacodontid found in Wyoming

The disappearance of the non-avian dinosaurs opened up an ecological niche for large mammalian herbivores. Some condylarths evolved to fill the niche, while others remained insectivorous. This may explain, in part, the tremendous evolutionary radiation of the condylarths that we can observe throughout the Paleocene, resulting in the different groups of ungulates (or "hoofed mammals") that form the dominant herbivores in most Cenozoic animal communities on land, except on the island continent of Australia.

Among recent mammals, Paenungulata (hyraxes, elephants, and sea cows), Perissodactyla (horses, rhinoceroses, and tapirs), Artiodactyla (pigs, deer, antelope, cows, camels, hippos, and their relatives), Cetacea (whales), and Tubulidentata (aardvarks) are traditionally regarded as members of the Euungulata.[9][18] Besides these, several extinct animals also belong to this group, especially the endemic South American orders of ungulates, (Meridiungulata). Although many ungulates have hooves, this feature does not define the Euungulata. Indeed, some condylarths had small hooves on their feet, but the most primitive forms are clawed.

Recent molecular and DNA research has reorganised the picture of mammalian evolution. Paenungulates and tubulidentates are seen as afrotherians, and no longer seen as closely related to the laurasiatherian perissodactyls, artiodactyls, and cetaceans,[19][20] implying that hooves were acquired independently (i.e. were analogous) by at least two different mammalian lineages, once in the Afrotheria and once in the Laurasiatheria. Condylarthra itself, therefore, is polyphyletic: the several condylarth groups are not closely related to each other at all. Indeed, Condylarthra is sometimes regarded as a 'wastebasket' taxon.[10] True relationships remain in many cases unresolved.

In addition to meridiungulates and living ungulates, a condylarthran ancestry has been proposed for several other extinct groups of mammals, including Mesonychia[7] and Dinocerata.[21]

See also

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Notes

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  1. ↑ Smith, De Bast. "Reassessment of the Small 'Arctocyonid' Prolatidens waudruae from the Early Paleocene of Belgium, and Its Phylogenetic Relationships with Ungulate-Like Mammals". Journal of Vertebrate Paleontology. Retrieved August 2013
  2. ↑ Ravel, Anthony; Orliac, Maeva (2014). "The inner ear morphology of the 'condylarthran' Hyopsodus lepidus". Historical Biology. 27 (8): 8. doi:10.1080/08912963.2014.915823. S2CID 84391276.
  3. ↑ Halliday, T.J.D.; Upchurch, P.; Goswami, A. (2017). "Resolving the relationships of Paleocene placental mammals". Biological Reviews. 92 (1): 521–550. doi:10.1111/brv.12242. PMC 6849585. PMID 28075073.
  4. ↑ Archibald, J. David; Zhang, Yue; Harper, Tony; Cifelli, Richard L. (May 6, 2011). "Protungulatum, confirmed Cretaceous occurrence of an otherwise Paleocene eutherian (placental?) mammal" (PDF). Journal of Mammalian Evolution. 18 (3): 153–161. doi:10.1007/s10914-011-9162-1. S2CID 16724836. Archived from the original (PDF) on March 3, 2016. Retrieved April 28, 2013.
  5. ↑ Cifelli, Richard L (1983-11-09). "The Origin and Affinities of the South American Condylarthra and Early Tertiary Litopterna (Mammalia)". American Museum Novitates (2772): 1–49.
  6. ↑ Bergqvist, Lilian P. (2008). "Postcranial Skeleton of the Upper Paleocene (Itaboraian) "Condylarthra" (Mammalia) of Itaboraí Basin, Brazil". In Sargis, Eric J.; Dagosto, Marian (eds.). Mammalian Evolutionary Morphology. Vertebrate Paleobiology and Paleoanthropology Series. Dordrecht: Springer Netherlands. pp. 107–133. doi:10.1007/978-1-4020-6997-0_6. ISBN 978-1-4020-6996-3.
  7. 1 2 Van Valen 1966
  8. 1 2 Rose, Kenneth D. (2006). "Archaic Ungulates". The beginning of the Age of Mammals. Baltimore: Johns Hopkins University Press. ISBN 978-0-8018-9221-9.
  9. 1 2 McKenna & Bell 1997
  10. 1 2 3 Janis 1993
  11. ↑ Naish, Darren (8 August 2013). "Phenacodontidae, I feel like I know you". Tetrapod Zoology. Scientific American. Archived from the original on 10 March 2014.
  12. ↑ Cooper, L. N.; Seiffert, E. R.; Clementz, M.; Madar, S. I.; Bajpai, S.; Hussain, S. T.; Thewissen, J. G. M. (2014-10-08). "Anthracobunids from the Middle Eocene of India and Pakistan Are Stem Perissodactyls". PLOS ONE. 9 (10) e109232. Bibcode:2014PLoSO...9j9232C. doi:10.1371/journal.pone.0109232. PMC 4189980. PMID 25295875.
  13. ↑ Chimento, Nicolás R.; Agnolin, Federico L. (2020). "Phylogenetic tree of Litopterna and Perissodactyla indicates a complex early history of hoofed mammals". Scientific Reports. 10 (1): 13280. Bibcode:2020NatSR..1013280C. doi:10.1038/s41598-020-70287-5. hdl:11336/13573. ISSN 2045-2322. PMC 7413542. PMID 32764723.
  14. ↑ James David Archibald · Alexander Olegovich Averianov, Phylogenetic analysis, taxonomic revision, and dental ontogeny of the Cretaceous Zhelestidae (Mammalia: Eutheria), Article · Feb 2012 · Zoological Journal of the Linnean Society
  15. ↑ Bast, Eric De; Smith, Thierry (2013-07-01). "Reassessment of the small 'arctocyonid' Prolatidens waudruae from the early Paleocene of Belgium, and its phylogenetic relationships with ungulate-like mammals". Journal of Vertebrate Paleontology. 33 (4): 964–976. Bibcode:2013JVPal..33..964B. doi:10.1080/02724634.2013.747531. ISSN 0272-4634.
  16. ↑ Halliday, Thomas J.D.; Upchurch, Paul; Goswami, Anjali (2015). "Resolving the relationships of Paleocene placental mammals" (PDF). Biological Reviews. 92 (1): 521–55. doi:10.1111/brv.12242. PMC 6849585. PMID 28075073. Archived (PDF) from the original on December 23, 2022.
  17. 1 2 Thewissen 1990, p. 20
  18. ↑ Novacek 1986
  19. ↑ Madsen et al. 2001
  20. ↑ Murphy et al. 2001
  21. ↑ Van Valen 1988

References

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  • Cope, E. D. (1881). "A new type of Perissodactyla". American Naturalist. 15 (12): 1017–20. doi:10.1086/272983. OCLC 45953517.
  • Cope, E. D. (1885). "The Vertebrata of the Tertiary Formations of the West". U. S. Geological Survey of the Territories. 3 (9): 1–1009. doi:10.1017/s0016756800467208. OCLC 3934701. Retrieved April 30, 2013.
  • Janis, C.M. (1993). "Tertiary Mammal Evolution in the Context of Changing Climates, Vegetation, and Tectonic Events". Annual Review of Ecology and Systematics. 24: 467–500. doi:10.1146/annurev.es.24.110193.002343.
  • Madsen, O.; Scally, M.; Douady, C.J.; Kao, D.; DeBry, R.W.; Adkins, R.; Amrine, H.M.; Stanhope, M.J.; de Jong, W.W.; Springer, M.S. (2001). "Parallel adaptive radiations in two major clades of placental mammals". Nature. 409 (6820): 610–614. Bibcode:2001Natur.409..610M. doi:10.1038/35054544. PMID 11214318. S2CID 4398233.
  • McKenna, M.C.; Bell, S.K. (1997). Classification of Mammals Above the Species Level. Columbia University Press. ISBN 978-0-231-11012-9.
  • Murphy, W.J.; Eizirik, E.; O'Brien, S.J.; Madsen, O.; Scally, M.; Douady, C.J.; Teeling, E.C.; Ryder, O.A.; Stanhope, M.J.; de Jong, W.W.; Springer, M.S. (2001). "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.
  • Novacek, M.J. (1986). "The skull of leptictid insectivorans and the higher-level classification of eutherian mammals". Bulletin of the American Museum of Natural History. 183 (1): 1–111. hdl:2246/1628.
  • Prothero, D.R.; Manning, E.M.; Fischer, M. (1988). "The phylogeny of the ungulates" (PDF). In Benton, M. J. (ed.). The phylogeny and classification of the tetrapods. 2: mammals. Systematics Association Special Volume 35B. Oxford: Clarendon Press. pp. 201–234. ISBN 978-0-19-857712-6. Retrieved May 1, 2013.
  • Thewissen, J.G.M. (1990). Evolution of Paleocene and Eocene Phenacodontidae (Mammalia, Condylarthra). Papers on Paleontology. Vol. 29. Museum of Paleontology, The University of Michigan. hdl:2027.42/48629. OCLC 742731818.
  • Van Valen, L.M. (1966). "Deltatheridia, a new order of mammals". Bulletin of the American Museum of Natural History. 132 (1): 1–126. hdl:2246/1126.
  • Van Valen, L.M. (1988). "Paleocene dinosaurs or Cretaceous ungulates in South America?". Evolutionary Monographs. 10: 1–79.
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