, Volume 98, Issue 12, pp 1027–1034 | Cite as

The last “pelycosaur”: a varanopid synapsid from the Pristerognathus Assemblage Zone, Middle Permian of South Africa

  • Sean P. Modesto
  • Roger M. H. Smith
  • Nicolás E. Campione
  • Robert R. ReiszEmail author
Original Paper


We report on a partial varanopid skull and mandible from the Pristerognathus Assemblage Zone of the Beaufort Group, in the South African Karoo Basin, which is probably latest Middle Permian (Capitanian) in age. This mycterosaurine is not only the youngest known varanopid from the Southern Hemisphere, but it is also the youngest known “pelycosaur” (i.e., non-therapsid synapsid). Like all other members of this clade of hypercarnivores, the teeth are strongly flattened, recurved, and have finely serrated cutting edges. The anterior dentary teeth form a caniniform region, and the splenial features a foramen intermandibularis oralis, the first ever to be described in a “pelycosaur.” The last varanopids were the smallest carnivores of latest Middle Permian continental faunas. Occupation of the small carnivore guild appears to have allowed varanopids to achieve a nearly cosmopolitan distribution throughout the Middle Permian, between the great Early Permian radiation of basal synapsids and the spectacular diversification of therapsid synapsids in the Late Permian and Early Triassic.


Paleozoic tetrapods Synapsida Varanopidae Paleobiogeography South Africa 



We thank Hedi Stummer and Annelise Crean for initial preparation of SAM-PK-K10407; Diane Scott for the photo images that appear in the Figures and Online Resource 1 of the Electronic supplementary material, and Nicola Wong Ken for additional preparation and the illustration of SAM-PK-K10407 and for the skull reconstruction seen in Fig. 3. Sheena Kaal is also thanked for supplemental locality data. This research was supported by Discovery Grants from the Natural Sciences and Engineering Research Council of Canada (to SPM and RRR).

Supplementary material

114_2011_856_MOESM1_ESM.doc (3.8 mb)
ESM 1 (DOC 3916 kb)


  1. Amson E, Laurin M (2011) On the affinities of Tetraceratops insignis, an Early Permian synapsid. Acta Palaeont Pol 56:301–312CrossRefGoogle Scholar
  2. Angielczyk KD, Rubidge BS (2009) The Permian dicynodont Colobodectes cluveri (Therapsida, Anomodontia), with notes on its ontogeny and stratigraphic range in the Karoo Basin, South Africa. J Vertebr Paleontol 29:1162–1173CrossRefGoogle Scholar
  3. Berman DS, Reisz RR (1982) Restudy of Mycterosaurus longiceps (Reptilia, Pelycosauria) from the Lower Permian of Texas. Ann Carnegie Mus 51:423–453Google Scholar
  4. Bond DPG, Hilton J, Wignall PB, Ali JR, Stevens LG, Sun Y, Lai X (2010) The Middle Permian (Capitanian) mass extinction on land and in the oceans. Earth Sci Rev 102:100–116CrossRefGoogle Scholar
  5. Botha-Brink J, Modesto SP (2007) A mixed-age classed ‘pelycosaur’ aggregation from South Africa: earliest evidence of parental care in amniotes? Proc R Soc B 274:2829–2834PubMedCrossRefGoogle Scholar
  6. Botha-Brink J, Modesto SP (2009) Anatomy and relationships of the Middle Permian varanopid Heleosaurus scholtzi based on a social aggregation from the Karoo Basin of South Africa. J Vertebr Paleontol 29:389–400CrossRefGoogle Scholar
  7. Botha-Brink J, Smith RMH (2006) Rapid vertebrate recuperation in the Karoo Basin of South Africa following the end-Permian extinction. J Afr Earth Sci 45:502–514CrossRefGoogle Scholar
  8. Campione NE, Reisz RR (2010) Varanops brevirostris (Eupelycosauria: Varanopidae) from the Lower Permian of Texas, with a discussion of varanopid morphology and interrelationships. J Vertebr Paleontol 30:724–746CrossRefGoogle Scholar
  9. Campione NE, Reisz RR (2011) Morphology and evolutionary significance of the atlas–axis complex in varanopid synapsids. Acta Palaeont Pol. doi: 10.4202/app. 2010.0071
  10. Dias-da-Silva S, Modesto SP, Schultz CL (2007) New material of Procolophon (Parareptilia: Procolophonoidea) from the Lower Triassic of Brazil, with remarks on the ages of the Sanga do Cabral and Buena Vista formations of South America. Can J Earth Sci 43:1685–1693CrossRefGoogle Scholar
  11. Dilkes DW, Reisz RR (1996) First record of a basal synapsid (‘mammal-like reptile’ in Gondwana. Proc R Soc B 263:1165–1170CrossRefGoogle Scholar
  12. Evans DC, Maddin HC, Reisz RR (2009) A re-evaluation of sphenacodontid synapsid material frm the Early Permian fissure fills at Richards Spur, Oklahoma. Palaeontology 52:219–227CrossRefGoogle Scholar
  13. Golubev VK (2005) Permian tetrapod stratigraphy. In: Lucas SG, Zeigler KE (eds) The nonmarine Permian. New Mexico Mus Nat Hist and Sci Bull 30:95–99Google Scholar
  14. Gradstein FM, Ogg JG (2004) Geologic time scale 2004 – why, how, and where next! Lethaia 37:175–181CrossRefGoogle Scholar
  15. Heaton MJ (1979) Cranial anatomy of primitive captorhinid reptiles from the Late Pennsylvanian and Early Permian [of] Oklahoma and Texas. Okla Geol Bull 127:1–84Google Scholar
  16. Ho Tun E (1979) Volcaniclastic material in the Lower Beaufort Group, Karoo rocks. Abstracts of the Geological Society of South Africa Geokongress ’79. Port Elizabeth 1:197–199Google Scholar
  17. Huttenlocker AK, Sidor CA, Smith RMH (2011) A new specimen of Promoschorhynchus (Therapsida: Therocephalia: Akidnognathidae) from the Lower Triassic of South Africa and its implications for theriodont survivorship across the Permo-Triassic boundary. J Vertebr Paleontol 31:405–421CrossRefGoogle Scholar
  18. Isozaki Y (2009) Illawarra Reversal: the fingerprint of a superplume that triggered Pangean breakup and the end-Guadalupian (Permian) mass extinction. Gond Res 15:421–432CrossRefGoogle Scholar
  19. Ivakhnenko MF (1991) Late Paleozoic tetrapod faunal assemblage from the Mezen River Basin. Paleont J 25:76–84Google Scholar
  20. Ivakhnenko MF (2008) Cranial morphology and evolution of Permian Dinomorpha (Eotherapsida) of Eastern Europe. Paleont J 42:859–995CrossRefGoogle Scholar
  21. Kemp TS (1982) Mammal-like reptiles and the origin of mammals. Academic, LondonGoogle Scholar
  22. Kutty TS (1972) Permian reptile fauna from India. Nature 237:462–463CrossRefGoogle Scholar
  23. Langston W Jr, Reisz RR (1981) Aerosaurus wellesi, new species, a varanopseid mammal-like reptile (Synapsida: Pelycosauria) from the Lower Permian of New Mexico. J Vertebr Paleontol 1:73–96CrossRefGoogle Scholar
  24. Laurin M, Reisz RR (1990) Tetraceratops is the oldest known therapsid. Nature 345:249–250CrossRefGoogle Scholar
  25. Laurin M, Reisz RR (1996) The osteology and relationships of Tetraceratops, the oldest known therapsid. J Vertebr Paleontol 16:95–102CrossRefGoogle Scholar
  26. Lucas SG (2006) Global Permian tetrapod biostratigraphy and biochronology. In: Lucas SG, Cassinis G, Schneider JW (eds) Non-marine Permian biostratigraphy and biochronology. Geol Soc Lond Spec Pub 265:65–93CrossRefGoogle Scholar
  27. Maddin HC, Sidor CA, Reisz RR (2008) Cranal anatomy of Ennatosaurus tecton (Synapsida: Caseidae) from the Middle Permian of Russia and the evolutionary relationships of Caseidae. J Vertebr Paleontol 28:160–180CrossRefGoogle Scholar
  28. Modesto SP, Rybczynski N (2000) The amniote faunas of the Russian Permian: implications for Late Permian terrestrial vertebrate biogeography. In: Benton MJ, Shishkin MA, Unwin DM, Kurochkin EN (eds) The age of dinosaurs in Russia and Mongolia. Cambridge University, Cambridge, pp 17–34Google Scholar
  29. Modesto SP, Smith RMH (2001) A new Late Permian captorhinid reptile: a first record from the South Africa Karoo. J Vert Palaeont 21:405–409CrossRefGoogle Scholar
  30. Modesto SP, Sidor CA, Rubidge BS, Welman J (2001) A second varanopseid skull from the Upper Permian of South Africa: implications for Late Permian ‘pelycosaur’ evolution. Lethaia 34:249–259CrossRefGoogle Scholar
  31. Nicolas M, Rubidge BS (2010) Changes in Permo-Triassic terrestrial tetrapod ecological representation in the Beaufort Group (Karoo Supergroup) of South Africa. Lethaia 43:45–59CrossRefGoogle Scholar
  32. O’Keefe FR, Sidor CA, Larsson HCE, Maga A, Ide O (2005) The vertebrate fauna of the Upper Permian of Niger-III, morphology and ontogeny of the hindlimb of Moradisaurus grandis (Reptilia, Captorhinidae). J Vert Paleont 25:309–319CrossRefGoogle Scholar
  33. Osborn HF (1903) On the primary division of the Reptilia into two sub-classes, Synapsida and Diapsida. Science 17:275–276PubMedCrossRefGoogle Scholar
  34. Pianka ER, King DR, King RA (eds) (2004) Varanoid lizards of the world. Indiana University, BloomingtonGoogle Scholar
  35. Piñeiro G, Verde M, Ubilla M, Ferigolo J (2003) First basal synapsids (“pelycosaurs”) from the Upper Permian-Lower Triassic of Uruguay, South America. J Paleont 77:389–392CrossRefGoogle Scholar
  36. Reisz RR (1986) Pelycosauria. In: Wellnhofer P (ed) Handbuch der Paläoherpetologie, Teil 17A. Gustav Fischer Verlag, pp 1–102Google Scholar
  37. Reisz RR, Berman DS (2001) The skull of Mesenosaurus romeri, a small varanopseid (Synapsida: Eupelycosauria) from the Upper Permian of the Mezen River Basin, northern Russia. Ann Carnegie Mus 70:113–132Google Scholar
  38. Reisz RR, Dilkes DW (2003) Archaeovenator hamiltonensis, a new varanopid (Synapsida: Eupelycosauria) from the Upper Carboniferous of Kansas. Can J Earth Sci 40:667–678CrossRefGoogle Scholar
  39. Reisz RR, Modesto SP (2007) Heleosaurus scholtzi from the Permian of South Africa: a varanopid synapsid, not a diapsid reptile. J Vertebr Paleontol 27:734–739CrossRefGoogle Scholar
  40. Reisz RR, Dilkes DW, Berman DS (1998) Anatomy and relationships of Elliotsmithia longiceps Broom, a small synapsid (Eupelycosauria: Varanopseidae) from the Late Permian of South Africa. J Vertebr Paleontol 18:602–611CrossRefGoogle Scholar
  41. Reisz RR, Liu J, Li J-L, Müller J (2011) A new captorhinid reptile, Gansurhinus qingtoushanensis, gen. et sp. nov., from the Permian of China. Naturwissenschaften 98:435–441PubMedCrossRefGoogle Scholar
  42. Retallack GJ, Metzger CA, Greaver T, Jahren AH, Smith RMH, Sheldon ND (2006) Middle-Late Permian mass extinction on land. GSA Bull 118:1398–1411CrossRefGoogle Scholar
  43. Romer AS (1966) Vertebrate paleontology. University of Chicago, ChicagoGoogle Scholar
  44. Romer AS, Price LI (1940) Review of the Pelycosauria. Geol Soc Amer Sp Pap 28:1–538Google Scholar
  45. Roussouw PJ, De Villiers J (1952) The geology of the Merweville area, Cape Province. Geological Survey of South Africa. Explanation to sheet 189Google Scholar
  46. Rubidge BS (1995) Biostratigraphy of the Eodicynodon Assemblage Zone. In: Rubidge BS (ed) Biostratigraphy of the Beaufort Group (Karoo Supergroup). S Afr Comm Strat, Pretoria, pp 3–7Google Scholar
  47. Rubidge BS (2005) Re-uniting lost continents—fossil reptiles from the ancient Karoo and their wanderlust. S Afr J Geol 108:135–172CrossRefGoogle Scholar
  48. Rubidge BS, Modesto S, Sidor CA, Welman J (1999) Eunotosaurus africanus from the Ecca-Beaufort contact in Northern Cape Province, South Africa—implications for Karoo Basin development. S Afr J Sci 95:553–555Google Scholar
  49. Rubidge BS, Sidor CA, Modesto SP (2006) A new burnetiamorph (Therapsida: Biarmosuchia) from the Middle Permian of South Africa. J Paleont 80:740–749CrossRefGoogle Scholar
  50. Smith RMH (1993) Vertebrate taphonomy of Late Permian floodplain deposits in the southwestern Karoo Basin of South Africa. Palaios 8:45–67CrossRefGoogle Scholar
  51. Smith RMH, Botha J (2005) The recovery of terrestrial vertebrate diversity in the South Africa Karoo Basin after the end-Permian extinction. C R Palevol 4:555–568CrossRefGoogle Scholar
  52. Smith RMH, Keyser AW (1995a) Biostratigraphy of the Tapinocephalus Assemblage Zone. In: Rubidge BS (ed) Biostratigraphy of the Beaufort Group (Karoo Supergroup). S Afr Comm Strat, Pretoria, pp 8–12Google Scholar
  53. Smith RMH, Keyser AW (1995b) Biostratigraphy of the Pristerognathus Assemblage Zone. In: Rubidge BS (ed) Biostratigraphy of the Beaufort Group (Karoo Supergroup). S Afr Comm Strat, Pretoria, pp 13–17Google Scholar
  54. Smith RMH, Ward PD (2001) Pattern of vertebrate extinctions across an event bed at the Permian-Triassic boundary in the Karoo Basin of South Africa. Geology 29:1147–1150CrossRefGoogle Scholar
  55. Smith RMH, Rubidge BS, Sidor CA (2006) A new burnetiid (Therapsida: Biarmosuchia) from the Upper Permian of South Africa and its biogeographic implications. J Vertebr Paleontol 26:331–343CrossRefGoogle Scholar
  56. Thulborn T, Turner S (2003) The last dicynodont: an Australian Cretaceous relict. Proc R Soc B 270:985–993PubMedCrossRefGoogle Scholar
  57. Welman J, Loock JC, Rubidge BS (2001) New evidence for diachroneity of the Ecca–Beaufort contact (Karoo Supergroup, South Africa). S Afr J Sci 97:320–322Google Scholar
  58. Williston SW (1914) The osteology of some American Permian vertebrates, I. Contrib Walker Mus 1:107–162Google Scholar

Copyright information

© Springer-Verlag 2011

Authors and Affiliations

  • Sean P. Modesto
    • 1
  • Roger M. H. Smith
    • 2
  • Nicolás E. Campione
    • 3
  • Robert R. Reisz
    • 4
    Email author
  1. 1.Department of BiologyCape Breton UniversitySydneyCanada
  2. 2.Karoo PalaeontologyIziko: South African MuseumCape TownSouth Africa
  3. 3.Department of Ecology and Evolutionary BiologyUniversity of TorontoTorontoCanada
  4. 4.Department of BiologyUniversity of Toronto at MississaugaMississaugaCanada

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