(Redirected from Pareiasauroidea)

Pareiasaurs (meaning "cheek lizards") are an extinct clade of large, herbivorous parareptiles. Members of the group were armoured with scutes which covered large areas of the body. They first appeared in southern Pangea during the Middle Permian, before becoming globally distributed during the Late Permian. Pareiasaurs were the largest reptiles of the Permian, reaching sizes equivalent to those of contemporary therapsids. Pareiasaurs became extinct at the end of the Permian during the Permian-Triassic extinction event.

Temporal range: Middle Permian - Late Permian, 265–252 Ma
Skeleton of Scutosaurus karpinskii in the American Museum of Natural History
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Parareptilia
Order: Procolophonomorpha
Node: Ankyramorpha
Suborder: Procolophonia
Clade: Pareiasauromorpha
Superfamily: Pareiasauroidea
Clade: Pareiasauria
Seeley, 1888


Restoration of Bradysaurus

Pareiasaurs ranged in size from 60 to 300 centimetres (2.0 to 9.8 ft) long, and may have weighed up to 600 kilograms (1,300 lb). They were stocky, with short tails, small heads, robust limbs, and broad feet. The cow-sized species Bunostegos, which lived 260 million years ago, is the earliest known example of a tetrapod with a fully erect posture as its legs were positioned directly under its body.[1] Pareiasaurs were protected by bony scutes called osteoderms that were set into the skin. Their heavy skulls were ornamented with multiple knobs and ridges. The leaf-shaped multi-cusped teeth resemble those of iguanas, caseids, and other reptilian herbivores. This dentition, together with the deep body, which may have housed an extensive digestive tract, are evidence of a herbivorous diet. Most authors have assumed a terrestrial lifestyle for pareiasaurs. A 2008 bone microanatomy study suggested a more aquatic, plausibly amphibious lifestyle,[2] but a later 2019 study found that the bone histology provided no direct evidence of this lifestyle.[3]

Evolutionary historyEdit

Pareiasaurs appear very suddenly in the fossil record. It is clear that these animals are parareptiles.[4][5] As such, they are closely related to Nycteroleterids.[6] Pareiasaurs filled the large herbivore niche (or guild) that had been occupied early in the Permian period by the Caseid pelycosaurs and, before them, the Diadectid reptillomorphs.[3] They are much larger than the diadectids, more similar to the giant caseid pelycosaur Cotylorhynchus. Although the last Pareiasaurs were no larger than the first types (indeed, many of the last ones became smaller), there was a definite tendency towards increased armour as the group developed. Pareiasaurs first appeared in the fossil record in the Middle Permian (Guadalupian) of Southern Pangaea, before dispersing into Northern Pangaea and gaining a cosmopolitan distribution during the Late Permian (Lopingian).[7]


Some paleontologists considered that pareiasaurs were direct ancestors of modern turtles. Pareiasaur skulls have several turtle-like features, and in some species the scutes have developed into bony plates, possibly the precursors of a turtle shell.[8] Jalil and Janvier, in a large analysis of pareiasaur relationships, also found turtles to be close relatives of the "dwarf" pareiasaurs, such as Pumiliopareia.[9] However, the discovery of Pappochelys argues against a potential pareisaurian relationship to turtles,[10] and DNA evidence indicates that living turtles are more closely related to living archosaurs than lepidosaurs, and therefore cladistically diapsids.[11]

Associated cladesEdit

Hallucicrania (Lee 1995): This clade was coined by MSY Lee for Lanthanosuchidae + (Pareiasauridae + Testudines). Lee's pareiasaur hypothesis has become untenable due to the diapsid features of the stem turtle Pappochelys and the potential testudinatan nature of Eunotosaurus. Recent cladistic analyses reveal that lanthanosuchids have a much more basal position in the Procolophonomorpha, and that the nearest sister taxon to the pareiasaurs are the rather unexceptional and conventional looking nycteroleterids (Müller & Tsuji 2007, Lyson et al. 2010) the two being united in the clade Pareiasauromorpha (Tsuji et al. 2012).

Pareiasauroidea (Nopcsa, 1928): This clade (as opposed to the superfamily or suborder Pareiasauroidea) was used by Lee (1995) for Pareiasauridae + Sclerosaurus. More recent cladistic studies place Sclerosaurus in the procolophonid subfamily Leptopleuroninae (Cisneros 2006, Sues & Reisz 2008), which means the similarities with pareiasaurs are the result of convergences.

Pareiasauria (Seeley, 1988): If neither Lanthanosuchids or Testudines are included in the clade, the Pareiasauria only contains the monophyletic family Pareiasauridae. It's a traditional linnaean term.


Below is a cladogram from Tsuji et al. (2013):[12]


"Bradysaurus" seeleyi

Bradysaurus baini













Pareiasuchus peringueyi

Pareiasuchus nasicornis








  1. ^ Pre-reptile may be earliest known to walk upright on all fours
  2. ^ Kriloff, A.; Germain, D.; Canoville, A.; Vincent, P.; Sache, M.; Laurin, M. (2008). "Evolution of bone microanatomy of the tetrapod tibia and its use in palaeobiological inference". Journal of Evolutionary Biology. 21 (3): 807–826. doi:10.1111/j.1420-9101.2008.01512.x. PMID 18312321. S2CID 6102313.
  3. ^ a b Boitsova, Elizaveta A; Skutschas, Pavel P; Sennikov, Andrey G; Golubev, Valeriy K; Masuytin, Vladimir V; Masuytina, Olga A (2019-07-05). "Bone histology of two pareiasaurs from Russia (Deltavjatia rossica and Scutosaurus karpinskii) with implications for pareiasaurian palaeobiology". Biological Journal of the Linnean Society: blz094. doi:10.1093/biolinnean/blz094. ISSN 0024-4066.
  4. ^ Gauthier, J., Kluge, A.G. and Rowe, T. (1988). "The early evolution of the Amniota." Pp. 103–155 in Benton, M.J. (ed.), The phylogeny and classification of the tetrapods, Volume 1: amphibians, reptiles, birds. Oxford: Clarendon Press.
  5. ^ Laurin, M.; Reisz, R.R. (1995). "A reevaluation of early amniote phylogeny". Zoological Journal of the Linnean Society. 113 (2): 165–223. doi:10.1111/j.1096-3642.1995.tb00932.x.
  6. ^ LEE, M. S. Y. (1995). "Historical burden in systematics and the interrelationships of 'parareptiles'". Biological Reviews of the Cambridge Philosophical Society. 70 (3): 459–547. doi:10.1111/j.1469-185x.1995.tb01197.x. S2CID 85790423.
  7. ^ Olroyd, Savannah L.; Sidor, Christian A. (August 2017). "A review of the Guadalupian (Middle Permian) global tetrapod fossil record". Earth-Science Reviews. 171: 583–597. doi:10.1016/j.earscirev.2017.07.001.
  8. ^ Lee, M.S.Y. (1997). "Pareiasaur phylogeny and the origin of turtles". Zoological Journal of the Linnean Society. 120 (3): 197–280. doi:10.1111/j.1096-3642.1997.tb01279.x.
  9. ^ Jalil, N.-E.; Janvier, P. (2005). "Les pareiasaures (Amniota, Parareptilia) du Permien supérieur du Bassin d'Argana, Maroc". Geodiversitas. 27 (1): 35–132.
  10. ^ Schoch, Rainer R.; Sues, Hans-Dieter (2015). "A Middle Triassic stem-turtle and the evolution of the turtle body plan". Nature. 523 (7562): 584–587. doi:10.1038/nature14472. PMID 26106865. S2CID 205243837.
  11. ^ Crawford, Nicholas G.; Parham, James F.; Sellas, Anna B.; Faircloth, Brant C.; Glenn, Travis C.; Papenfuss, Theodore J.; Henderson, James B.; Hansen, Madison H.; Simison, W. Brian (February 2015). "A phylogenomic analysis of turtles". Molecular Phylogenetics and Evolution. 83: 250–257. doi:10.1016/j.ympev.2014.10.021. PMID 25450099.
  12. ^ Tsuji, L. A.; Sidor, C. A.; Steyer, J. - S. B.; Smith, R. M. H.; Tabor, N. J.; Ide, O. (2013). "The vertebrate fauna of the Upper Permian of Niger—VII. Cranial anatomy and relationships of Bunostegos akokanensis (Pareiasauria)". Journal of Vertebrate Paleontology. 33 (4): 747–763. doi:10.1080/02724634.2013.739537. S2CID 86097405.

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