Skip to main content

Permian and Triassic Dicynodont (Therapsida: Anomodontia) Faunas of the Luangwa Basin, Zambia: Taxonomic Update and Implications for Dicynodont Biogeography and Biostratigraphy

  • Chapter
  • First Online:
Early Evolutionary History of the Synapsida

Abstract

Dicynodont fossils were first collected in the Luangwa Basin, Zambia, in the 1920s, but limited detailed study and taxonomic uncertainty have obscured their biostratigraphic utility and their implications for topics such as dicynodont biogeography and the effects of the end-Permian extinction. Here we present a comprehensive taxonomic revision of the dicynodonts of the Luangwa Basin, taking into account specimens in all major museum collections and new material collected by our team in 2009. We recognize 14 dicynodont species from the Upper Permian Upper Madumabisa Mudstone: Pristerodon mackayi, Endothiodon sp., Diictodon feliceps, Compsodon helmoedi, Emydops sp., Dicynodontoides cf. D. nowacki, a new tusked cistecephalid, cf. Katumbia parringtoni, Kitchinganomodon crassus, Oudenodon bainii, Odontocyclops whaitsi, Dicynodon huenei, Syops vanhoepeni, and a new lystrosaurid. Previous reports of Lystrosaurus in the basin appear to be in error. In addition, we found no significant partitioning of dicynodont taxa in the northern and southern parts of the basin, despite substantial differences in preservation, indicating the presence of a single faunal assemblage in the Upper Permian. The Madumabisa dicynodont assemblage is best correlated with the Cistecephalus Assemblage Zone of South Africa. The shared presence of Dicynodon huenei and possibly Katumbia in the Luangwa Basin and the Ruhuhu Basin of Tanzania suggests that the Tanzanian Usili Formation also can be correlated with the Cistecephalus zone. Interestingly, the Madumabisa assemblage from Zambia is more similar to the coeval assemblage from South Africa, despite its closer geographic proximity to Tanzania. The Karoo and Ruhuhu basins also include more endemic species in the Permian than the Luangwa Basin. The Middle Triassic Ntawere Formation preserves four dicynodont species (Kannemeyeria lophorhinus, “Kannemeyerialatirostris, Zambiasaurus submersus, Sangusaurus edentatus), which occur at two stratigraphic levels. The lower Ntawere assemblage resembles that of the Omingonde Formation of Namibia in the presence of Kannemeyeria lophorhinus and potentially Dolichuranus (if “K.latirostris represents this taxon). The upper Ntawere assemblage shares the genus Sangusaurus with that of the Manda beds of Tanzania and includes the endemic Zambiasaurus. Comparisons of these assemblages to the Omingonde and Manda suggest that both are best correlated with the Cynognathus C subzone. When combined with data on other tetrapod taxa, our revised dicynodont assemblages contribute to an emerging picture of broad faunal similarity in southern and eastern Africa during the Late Permian, and increasing differentiation between the South African and other Karoo basins following the end-Permian extinction.

This is a preview of subscription content, log in via an institution to check access.

Access this chapter

Chapter
USD 29.95
Price excludes VAT (USA)
  • Available as PDF
  • Read on any device
  • Instant download
  • Own it forever
eBook
USD 99.00
Price excludes VAT (USA)
  • Available as EPUB and PDF
  • Read on any device
  • Instant download
  • Own it forever
Softcover Book
USD 129.00
Price excludes VAT (USA)
  • Compact, lightweight edition
  • Dispatched in 3 to 5 business days
  • Free shipping worldwide - see info
Hardcover Book
USD 129.00
Price excludes VAT (USA)
  • Durable hardcover edition
  • Dispatched in 3 to 5 business days
  • Free shipping worldwide - see info

Tax calculation will be finalised at checkout

Purchases are for personal use only

Institutional subscriptions

References

  • Abdala, F., & Allinson, M. (2005). The taxonomic status of Parathrinaxodon proops (Therapsida: Cynodontia), with comments on the morphology of the palate in basal cynodonts. Palaeontologia Africana, 41, 45–52.

    Google Scholar 

  • Abdala, F., Hancox, P. J., & Neveling, J. (2005). Cynodonts from the uppermost Burgersdorp Formation, South Africa, and their bearing on the biostratigraphy and correlation of the Triassic Cynognathus Assemblage Zone. Journal of Vertebrate Paleontology, 25, 192–199.

    Google Scholar 

  • Anderson, J. M., & Cruickshank, A. R. I. (1978). The biostratigraphy of the Permian and the Triassic. Part 5. A review of the classification and distribution of Permo-Triassic tetrapods. Palaeontologia Africana, 21, 15–44.

    Google Scholar 

  • Angielczyk, K. D. (2002). Redescription, phylogenetic position, and stratigraphic significance of the dicynodont genus Odontocyclops (Synapsida: Anomodontia). Journal of Paleontology, 76, 1047–1059.

    Google Scholar 

  • Angielczyk, K. D. (2007). New specimens of the Tanzanian dicynodont “Cryptocynodonparringtoni von Huene, 1942 (Therapsida, Anomodontia), with an expanded analysis of Permian dicynodont phylogeny. Journal of Vertebrate Paleontology, 27, 116–131.

    Google Scholar 

  • Angielczyk, K. D., & Kurkin, A. A. (2003). Phylogenetic analysis of Russian Permian dicynodonts (Therapsida: Anomodontia): Implications for Permian biostratigraphy and Pangaean biogeography. Zoological Journal of the Linnean Society, 139, 157–212.

    Google Scholar 

  • Angielczyk, K. D., & Rubidge, B. S. (2010). A new pylaecephalid dicynodont (Therapsida, Anomodontia) from the Tapinocephalus Assemblage Zone, Karoo Basin, Middle Permian of South Africa. Journal of Vertebrate Paleontology, 30, 1396–1409.

    Google Scholar 

  • Angielczyk, K. D., & Sullivan, C. (2008). Diictodon feliceps (Owen, 1876), a dicynodont (Therapsida, Anomodontia) species with a Pangaean distribution. Journal of Vertebrate Paleontology, 28, 788–802.

    Google Scholar 

  • Angielczyk, K. D., Fröbisch, J., & Smith, R. M. H. (2005). On the stratigraphic range of the dicynodont taxon Emydops (Therapsida: Anomodontia) in the Karoo Basin, South Africa. Palaeontologia Africana, 41, 23–33.

    Google Scholar 

  • Angielczyk, K. D., Sidor, C. A., Nesbitt, S. J., Smith, R. M. H., & Tsuji, L. A. (2009). Taxonomic revision and new observations on the postcranial skeleton, biogeography, and biostratigraphy of the dicynodont genus Dicynodontoides, the senior subjective synonym of Kingoria (Therapsida, Anomodontia). Journal of Vertebrate Paleontology, 29, 1174–1187.

    Google Scholar 

  • Angielczyk, K. D., & Rubidge, B. S. (2013). Skeletal morphology, phylogenetic relationships, and stratigraphic range of Eosimops newtoni Broom, 1921, a pylaecephalid dicynodont (Therapsida, Anomodontia) from the Middle Permian of South Africa. Journal of Systematic Palaeontology, 11, 191–231.

    Google Scholar 

  • Attridge, J., Ball, H. W., Charig, A. J., & Cox, C. B. (1964). The British Museum (Natural History)—University of London joint palaeontological expedition to Northern Rhodesia and Tanganyika. Nature, 201, 445–449.

    Google Scholar 

  • Bandyopadhyay, S. (1985). Dicynodont reptiles from the Triassic Yerrapalli Formation and their importance in stratigraphy and palaeontology. Unpublished Ph.D. thesis, Calcutta University, Calcutta.

    Google Scholar 

  • Bandyopadhyay, S. (1988). A kannemeyeriid dicynodont from the Middle Triassic Yerrapalli Formation. Philosophical Transactions of the Royal Society Series B, 320, 185–233.

    Google Scholar 

  • Bandyopadhyay, S. (1989). The mammal-like reptile Rechnisaurus from the Triassic of India. Palaeontology, 32, 305–312.

    Google Scholar 

  • Banks, N. L., Bardwell, K. A., & Musiwa, S. (1995). Karoo rift basins of the Luangwa Valley, Zambia. In J. J. Lambiase (Ed.), Hydrocarbon habitat in rift basins (Vol. 80, pp. 285–295). London: Geological Society, Special Publications

    Google Scholar 

  • Battail, B. (1978). Les Reptiles Thérapsides dans la stratigraphie du Trias continental: les faunes d’âge Stormberg en Afrique et leurs équivalents dans le monde. Société Géologique du Nord Annales, 97, 343–350.

    Google Scholar 

  • Battail, B. (1993). On the biostratigraphy of Triassic therapsid-bearing formations. In S. G. Lucas & M. Morales (Eds.), The nonmarine Triassic (pp. 31–35). New Mexico Museum of Natural History and Science Bulletin, 3.

    Google Scholar 

  • Battail, B. (2009a). Late Permian dicynodont fauna from Laos. In E. Buffetaut, G. Cuny, J. Le Loeuff, & V. Suteethorn (Eds.), Late Paleozoic and Mesozoic ecosystems in Southeast Asia (Vol. 315, pp. 33–40). London: Geological Society, Special Publications.

    Google Scholar 

  • Battail, B. (2009b). Dicynodon from South Africa, ‘Dicynodon’ from Russia, and Dicynodon(?) from southeast Asia. Palaeontologia Africana, 44, 143–145.

    Google Scholar 

  • Bond, G. (1967). A review of Karroo sedimentation and lithology in southern Rhodesia. In Reviews prepared for the First Symposium on Gondwana Stratigraphy (pp. 173–195). Mar del Plata: IUGS.

    Google Scholar 

  • Boonstra, L. D. (1938). A report on some Karroo reptiles from the Luangwa Valley, Northern Rhodesia. Quarterly Journal of the Geological Society of London, 94, 371–384.

    Google Scholar 

  • Botha, J., & Angielczyk, K. D. (2007). An integrative approach to distinguishing the Late Permian dicynodont species Oudenodon bainii and Tropidostoma microtrema (Therapsida, Anomodontia). Palaeontology, 50, 1175–1209.

    Google Scholar 

  • Botha, J., & Smith, R. M. H. (2006). Rapid vertebrate recuperation in the Karoo Basin of South Africa following the end-Permian extinction. Journal of African Earth Sciences, 45, 502–514.

    Google Scholar 

  • Botha, J., & Smith, R. M. H. (2007). Lystrosaurus species composition across the Permo-Triassic boundary in the Karoo Basin of South Africa. Lethaia, 40, 125–137.

    Google Scholar 

  • Botha-Brink, J., Huttenlocker, A. K., & Modesto, S. P. (2013). Vertebrate paleontology of Nooitgedacht 68: A Lystrosaurus maccaigi-rich Permo-Triassic boundary locality in South Africa. In C. F. Kammerer, K. D. Angielczyk, & J. Fröbisch (Eds.), Early evolutionary history of the Synapsida (pp. 289–304). Dordrecht: Springer.

    Google Scholar 

  • Brink, A. S. (1963). Two cynodonts from the N’tawere Formation in the Luangwa Valley of Northern Rhodesia. Palaeontologia Africana, 8, 77–96.

    Google Scholar 

  • Brink, A. S., & Keyser, A. W. (1983). Illustrated bibliographic catalogue of the Synapsida. Geological Survey of South Africa Handbook, 10, J212A231B1.

    Google Scholar 

  • Broom, R. (1903). On the classification of the theriodonts and their allies. Report of the South African Association for the Advancement of Science, 1, 286–294.

    Google Scholar 

  • Broom, R. (1905). On the use of the term Anomodontia. Albany Museum Records, 1, 266–269.

    Google Scholar 

  • Broom, R. (1913). On some new genera and species of dicynodont reptiles, with notes on a few others. American Museum of Natural History Bulletin, 32, 441–457.

    Google Scholar 

  • Broom, R. (1935). A new type of anomodont reptile. Nature, 135, 583–584.

    Google Scholar 

  • Cairncross, B. (2001). An overview of the Permian (Karoo) coal deposits of southern Africa. Journal of African Earth Sciences, 33, 529–562.

    Google Scholar 

  • Camp, C. A., & Welles, S. P. (1956). Triassic dicynodont reptiles. Part I. The North American genus Placerias. Memoirs of the Univeristy of California, 13, 255–304.

    Google Scholar 

  • Catuneanu, O., Wopfner, H., Eriksson, P. G., Cairncross, B., Rubidge, B. S., Smith, R. M. H., et al. (2005). The Karoo basins of south-central Africa. Journal of African Earth Sciences, 43, 211–253.

    Google Scholar 

  • Chernin, S. (1974). Capitosaurid amphibians from the Upper Luangwa Valley, Zambia. Palaeontologia Africana, 17, 33–55.

    Google Scholar 

  • Cluver, M. A. (1971). The cranial morphology of the dicynodont genus Lystrosaurus. Annals of the South African Museum, 56, 155–274.

    Google Scholar 

  • Cluver, M. A. (1974a). The skull and mandible of a new cistecephalid dicynodont. Annals of the South African Museum, 64, 137–155.

    Google Scholar 

  • Cluver, M. A. (1974b). The cranial morphology of the Lower Triassic dicynodont Myosaurus gracilis. Annals of the South African Museum, 66, 35–54.

    Google Scholar 

  • Cluver, M. A. (1978). The skeleton of the mammal-like reptile Cistecephalus with evidence for a fossorial mode of life. Annals of the South African Museum, 76, 213–246.

    Google Scholar 

  • Cluver, M. A., & Hotton, N. (1981). The genera Dicynodon and Diictodon and their bearing on the classification of the Dicynodontia (Reptilia, Therapsida). Annals of the South African Museum, 83, 99–146.

    Google Scholar 

  • Cluver, M. A., & King, G. M. (1983). A reassessment of the relationships of Permian Dicynodontia (Reptilia, Therapsida) and a new classification of dicynodonts. Annals of the South African Museum, 91, 195–273.

    Google Scholar 

  • Cooper, M. R. (1980). “The origins and classification of Triassic Dicynodonts” by A. W. Keyser and A. R. I. Cruickshank. Discussion. Transactions of the Geological Society of South Africa, 83, 107–110.

    Google Scholar 

  • Cooper, M. R. (1982). A mid-Permian to earliest Jurassic tetrapod biostratigraphy and its significance. Arnoldia Zimbabwe, 9, 77–104.

    Google Scholar 

  • Cope, E. D. (1871). On the homologies of some of the cranial bones of the Reptilia, and on the systematic arrangement of the class. Proceedings of the American Association for the Advancement of Science, 19, 194–247.

    Google Scholar 

  • Cosgriff, J. W., Hammer, W. R., & Ryan, W. J. (1982). The Pangaean reptile Lystrosaurus maccaigi in the Lower Triassic of Antarctica. Journal of Paleontology, 56, 371–385.

    Google Scholar 

  • Cox, C. B. (1964). On the palate, dentition, and classification of the fossil reptile Endothiodon and related genera. American Museum Novitates, 2171, 1–25.

    Google Scholar 

  • Cox, C. B. (1969). Two new dicynodonts from the Triassic Ntawere Formation, Zambia. Bulletin of the British Museum (Natural History), 17, 257–294.

    Google Scholar 

  • Cox, C. B. (1972). A new digging dicynodont from the Upper Permian of Tanzania. In K. A. Joysey & T. S. Kemp (Eds.), Studies in vertebrate evolution (pp. 173–189). Edinburgh: Oliver & Boyd.

    Google Scholar 

  • Cox, C. B. (1991). The Pangaea dicynodont Rechnisaurus and the comparative biostratigraphy of Triassic dicynodont faunas. Palaeontology, 34, 767–784.

    Google Scholar 

  • Cox, C. B. (1998). The jaw function and adaptive radiation of the dicynodont mammal-like reptiles of the Karoo Basin of South Africa. Zoological Journal of the Linnean Society, 122, 349–384.

    Google Scholar 

  • Cox, C. B., & Li, J.-L. (1983). A new genus of Triassic dicynodont from East Africa and its classification. Palaeontology, 26, 389–406.

    Google Scholar 

  • Crozier, E. A. (1970). Preliminary report on two Triassic dicynodonts form Zambia. Palaeontologia Africana, 13, 39–45.

    Google Scholar 

  • Cruickshank, A. R. I. (1967). A new dicynodont genus from the Manda Formation of Tanzania (Tanganyika). Journal of Zoology, 153, 163–208.

    Google Scholar 

  • Cruickshank, A. R. I. (1986). Biostratigraphy and classification of a new Triassic dicynodont from East Africa. Modern Geology, 10, 121–131.

    Google Scholar 

  • Cruickshank, A. R. I., Clark, N. D. L., & Adams, C. (2005). A new specimen of Dicynodon traquairi (Newton) (Synapsida: Anomodontia) from Elgin, Scotland. Palaeontologia Africana, 41, 35–43.

    Google Scholar 

  • Damiani, R., Vasconcelos, C., Renaut, A., Hancox, J., & Yates, A. (2007). Dolichuranus primaevus (Therapsida: Anomodontia) from the Middle Triassic of Namibia and its phylogenetic relationships. Palaeontology, 50, 1531–1546.

    Google Scholar 

  • Davies, K. C. (1981). Heulandite deposition in fossil reptile bones from the Karroo of Zambia. Journal of Paleontology, 55, 47–61.

    Google Scholar 

  • DeFauw, S. L. (1989). Patterns of evolution in the Dicynodontia, with special reference to austral taxa. In J. A. Crame (Ed.), Origins and evolution of the Antarctic Biota (Vol. 47, pp. 63–84). London: Geological Society, Special Publications.

    Google Scholar 

  • DeFauw, S. L. (1993). The Pangaean dicynodont Rechnisaurus from the Triassic of Argentina. In S. G. Lucas & M. Morales (Eds.), The nonmarine Triassic (pp. 101–105). New Mexico Museum of Natural History and Science Bulletin, 3.

    Google Scholar 

  • Dixey, F. (1937). The geology of part of the upper Luangwa Valley, north-eastern Rhodesia. Quarterly Journal of the Geological Society of London, 93, 52–74.

    Google Scholar 

  • Drysdall, A. R., & Kitching, J. W. (1962). The Karoo succession of the Upper Luangwa Valley, Northern Rhodesia. Transactions and Proceedings of the Geological Society of South Africa, 65, 75–90.

    Google Scholar 

  • Drysdall, A. R., & Kitching, J. W. (1963). A re-examination of the Karroo succession and fossil localities of part of the upper Luangwa Valley. Geological Survey of Northern Rhodesia Memoir, 1, 1–62.

    Google Scholar 

  • Freeman, L. (1993). The cranial morphology of a new tusked species of the genus Cistecephalus (Therapsida, Dicynodontia). Unpublished B.Sc. Honours thesis, University of the Witwatersrand, Johannesburg.

    Google Scholar 

  • Fröbisch, J. (2007). The cranial anatomy of Kombuisia frerensis Hotton (Synapsida, Dicynodontia) and a new phylogeny of anomodont therapsids. Zoological Journal of the Linnean Society, 150, 117–144.

    Google Scholar 

  • Fröbisch, J. (2008). Global taxonomic diversity of anomodonts (Tetrapoda, Therapsida) and the terrestrial rock record across the Permian-Triassic boundary. PLoS ONE, 3, e3733. doi: 10.1371/journal.pone.0003733.

    Google Scholar 

  • Fröbisch, J. (2009). Composition and similarity of global anomodont-bearing tetrapod faunas. Earth-Science Reviews, 95, 119–175.

    Google Scholar 

  • Fröbisch, J., & Reisz, R. R. (2008). A new species of Emydops (Synapsida, Anomodontia) and a discussion of dental variability and pathology in dicynodonts. Journal of Vertebrate Paleontology, 28, 770–787.

    Google Scholar 

  • Gair, S. H. (1959). The Karroo System and coal resources of the Gwembe District, north-east section. Geological Survey of Northern Rhodesia Bulletin, 1, 1–88.

    Google Scholar 

  • Gale, T. M. (1988). Comments on a “nest” of juvenile dicynodont reptiles. Modern Geology, 13, 119–124.

    Google Scholar 

  • Gale, T. M. (1989). Function, ecology, and relationships of fossil herbivores. Unpublished Ph.D. thesis, University of Oxford, Oxford.

    Google Scholar 

  • Gay, S. A., & Cruickshank, A. R. I. (1999). Biostratigraphy of the Permian tetrapod faunas from the Ruhuhu Valley, Tanzania. Journal of African Earth Sciences, 29, 195–210.

    Google Scholar 

  • Govender, R., & Yates, A. (2009). Dicynodont postcrania from the Triassic of Namibia and their implications for the systematics of Kannemeyeriiforme dicynodonts. Palaeontologia Africana, 44, 41–57.

    Google Scholar 

  • Govender, R., Hancox, P. J., & Yates, A. M. (2008). Re-evaluation of the postcranial skeleton of the Triassic dicynodont Kannemeyeria simocephalus form the Cynognathus Assemblage Zone (Subzone B) of South Africa. Palaeontologia Africana, 43, 19–37.

    Google Scholar 

  • Grine, F. E., Foster, C. A., Cluver, M. A., & Georgi, J. A. (2006). Cranial variability, ontogeny, and taxonomy of Lystrosaurus from the Karoo Basin of South Africa. In M. T. Carrano, T. J. Gaudin, R. W. Blob, & J. R. Wible (Eds.), Amniote paleobiology (pp. 432–503). Chicago: University of Chicago Press.

    Google Scholar 

  • Hancox, P. J. (2000). The continental Triassic of South Africa. Zentralblatt für Geologie und Paläontologie Teil I, 1998, 1285–1324.

    Google Scholar 

  • Hancox, P. J., & Rubidge, B. S. (1997). The role of fossils in interpreting the development of the Karoo Basin. Palaeontologia Africana, 33, 41–54.

    Google Scholar 

  • Hancox, P. J., & Rubidge, B. S. (2001). Breakthroughs in the biodiversity, biogeography, biostratigraphy, and basin analysis of the Beaufort Group. Journal of African Earth Sciences, 33, 563–577.

    Google Scholar 

  • Hancox, P. J., Angielczyk, K. D., & Rubidge B. S. (2013). Angonisaurus and Shansiodon, dicynodonts (Therapsida, Anomodontia) from subzone C of the Cynognathus Assemblage Zone (Middle Triassic) of South Africa. Journal of Vertebrate Paleontology, 33, 655–676.

    Google Scholar 

  • Haughton, S. H. (1926). On Karroo vertebrates from Nyasaland. Transactions of the Geological Society of South Africa, 27, 69–83.

    Google Scholar 

  • Haughton, S. H. (1932). On a collection of Karroo vertebrates from Tanganyika Territory. Quarterly Journal of the Geological Society of London, 88, 634–668.

    Google Scholar 

  • Huene, F. von. (1942). Die Anomodontier des Ruhuhu-Gebietes in der Tübinger Sammlung. Palaeontographica Abteilung A, 44, 154–184.

    Google Scholar 

  • Huene, F. von. (1948). Short review of the lower tetrapods. In A. Du Toit (Ed.), Robert Broom Commemorative Volume (pp. 65–106). Cape Town: Royal Society of South Africa.

    Google Scholar 

  • Huxley, T. H. (1868). On Saurosternon Bainii, and Pristerodon McKayi, two new fossil lacertillian reptiles from South Africa. Geological Magazine, 5, 201–205.

    Google Scholar 

  • Irmis, R. B. (2005). The vertebrate fauna of the Upper Triassic Chinle Formation in northern Arizona. In S. J. Nesbitt, W. G. Parker, & R. B. Irmis (Eds.), Guidebook to the Triassic Formations of the Colorado Plateau in Northern Arizona (pp. 63–88). Mesa Southwest Museum Bulletin, 9.

    Google Scholar 

  • Ivakhnenko, M. F., Golubev, V. K., Gubin, Y. M., Kalandadze, N. N., Novikov, I. V., Sennikov, A. G., et al. (1997). Permian and Triassic Tetrapods of Eastern Europe. Geos: Moscow.

    Google Scholar 

  • Jacobs, L. L., Winkler, D. A., Newman, K. D., Gomani, E. M., & Deino, A. (2005). Therapsids from the Late Permian Chiweta Beds and the age of the Karoo Supergroup in Malawi. Palaeontologia Electronica, 8, 1–23.

    Google Scholar 

  • Jain, S. L., & Roy Chowdhury, T. (1987). Fossil vertebrates from the Pranhita-Godavari Valley (India) and their stratigraphic correlation. In J. W. Collinson, D. H. Elliot, S. M. Haban, & G. D. McKenzie (Eds.), Gondwana six: Stratigraphy, sedimentology, and paleontology (pp. 219–228). Geophysical Monograph, 41.

    Google Scholar 

  • Kammerer, C. F., & Angielczyk, K. D. (2009). A proposed higher taxonomy of anomodont therapsids. Zootaxa, 2018, 1–24.

    Google Scholar 

  • Kammerer, C. F., Angielczyk, K. D., & Fröbisch, J. (2011). A comprehensive taxonomic revision of Dicynodon (Therapsida, Anomodontia), and its implications for dicynodont phylogeny, biogeography, and biostratigraphy. Society of Vertebrate Paleontology Memoir, 11, 1–158.

    Google Scholar 

  • Kemp, T. S. (1975). Vertebrate localities in the Karroo System of the Luangwa Valley, Zambia. Nature, 254, 415–416.

    Google Scholar 

  • Kemp, T. S. (1979). The primitive cynodont Procynosuchus: Functional anatomy of the skull and relationships. Philosophical Transactions of the Royal Society of London. Series B, 285, 73–122.

    Google Scholar 

  • Kemp, T. S. (1980a). Aspects of the structure and functional anatomy of the Middle Triassic cynodont Luangwa. Journal of Zoology, 191, 193–239.

    Google Scholar 

  • Kemp, T. S. (1980b). The primitive cynodont Procynosuchus: Structure, function and evolution of the postcranial skeleton. Philosophical Transactions of the Royal Society of London. Series B, 288, 217–258.

    Google Scholar 

  • Kemp, T. S. (2005). The origin and evolution of mammals. Oxford: Oxford University Press.

    Google Scholar 

  • Kerr, C. D. (1974). Preliminary report on the 1974 Geological Survey of Zambia & Oxford University Museum Joint Palaeontological Expedition to Luangwa Valley. Unpublished manuscript, Geological Survey of Zambia, Lusaka.

    Google Scholar 

  • Keyser, A. W. (1972). A re-evaluation of the systematics and morphology of certain anomodont Therapsida. Palaeontologia Africana, 14, 15–16.

    Google Scholar 

  • Keyser, A. W. (1973a). A re-evaluation of the genus Tropidostoma Seeley. Palaeontologia Africana, 16, 25–35.

    Google Scholar 

  • Keyser, A. W. (1973b). A preliminary study of the type area of the Cistecephalus Zone of the Beaufort Series, and a revision of the anomodont family Cistecephalidae. Geological Survey of South Africa Memoir, 62, 1–71.

    Google Scholar 

  • Keyser, A. W. (1973c). A new Triassic vertebrate fauna from South West Africa. Palaeontologia Africana, 16, 1–15.

    Google Scholar 

  • Keyser, A. W. (1973d). New Triassic vertebrate fauna from South West Africa. South African Journal of Science, 69, 113–115.

    Google Scholar 

  • Keyser, A. W. (1974). Evolutionary trends in Triassic Dicynodontia. Palaeontologia Africana, 17, 57–68.

    Google Scholar 

  • Keyser, A. W. (1975). A reevaluation of the cranial morphology and systematics of some tuskless Anomodontia. Geological Survey of South Africa Memoir, 67, 1–110.

    Google Scholar 

  • Keyser, A. W. (1979). A new dicynodont genus and its bearing on the origin of the Gondwana Triassic Dicynodontia. In B. Laskar & C. S. Raja Rao (Eds.), Proceedings and papers of the 4th IUGS Gondwana symposium (pp. 184–198). Delhi: Hindustan Publishing Corporation.

    Google Scholar 

  • Keyser, A. W. (1981). The stratigraphic distribution of the Dicynodontia of Africa reviewed in a Gondwana context. In M. M. Creswell & P. Vella (Eds.), Gondwana five (pp. 61–63). Rotterdam: A.A. Balkema.

    Google Scholar 

  • Keyser, A. W. (1993). A re-evaluation of the smaller Endothiodontidae. Geological Survey of South Africa Memoir, 82, 1–53.

    Google Scholar 

  • Keyser, A. W., & Cruickshank, A. R. I. (1979). The origins and classification of Triassic dicynodonts. Transactions of the Geological Society of South Africa, 82, 81–108.

    Google Scholar 

  • Keyser, A. W., & Cruickshank, A. R. I. (1980). “The origins and classification of Triassic Dicynodonts” by A. W. Keyser and A. R. I. Cruickshank. Author’s reply to discussion. Transactions of the Geological Society of South Africa, 83, 110–111.

    Google Scholar 

  • King, G. M. (1981). The functional anatomy of a Permian dicynodont. Philosophical Transactions of the Royal Society of London. Series B, 291, 243–322.

    Google Scholar 

  • King, G. M. (1988). Anomodontia. In P. Wellnhofer (Ed.), Handbuch der Paläoherpetologie (Vol. 17C). Stuttgart: Gustav Fischer Verlag.

    Google Scholar 

  • King, G. M. (1990). The dicynodonts: A study in palaeobiology. London: Chapman and Hall.

    Google Scholar 

  • King, G. M. (1992). The paleobiogeography of Permian anomodonts. Terra Nova, 4, 633–640.

    Google Scholar 

  • King, G. M. (1993). How many species of Diictodon were there? Annals of the South African Museum, 102, 303–325.

    Google Scholar 

  • King, G. M., & Jenkins, I. (1997). The dicynodont Lystrosaurus from the Upper Permian of Zambia: Evolutionary and stratigraphical implications. Palaeontology, 40, 149–156.

    Google Scholar 

  • King, G. M., & Rubidge, B. S. (1993). A taxonomic revision of small dicynodonts with postcanine teeth. Zoological Journal of the Linnean Society, 107, 131–154.

    Google Scholar 

  • Kitching, J. W. (1963). The fossil localities and mammal-like reptiles of the upper Luangwa Valley, Northern Rhodesia. South African Journal of Science, 59, 259–264.

    Google Scholar 

  • Kitching, J. W. (1977). The distribution of the Karroo vertebrate fauna. Bernard Price Institute for Palaeontological Research Memoir, 1, 1–131.

    Google Scholar 

  • Kutty, T. S. (1972). Permian reptilian fauna from India. Nature, 237, 462–463.

    Google Scholar 

  • Larkin, N. (1994). Description of a new Triassic dicynodont from the Manda Formation of Tanzania. Unpublished M.Sc. thesis, University College London, London.

    Google Scholar 

  • Lehman J.-P. (1961). Dicynodontia. In Piveteau, J. (Ed.), Traité de Paléontologie. VI(i) Mammifères. Origine Reptilienne. Evolution (pp. 287–351). Paris: Masson et Cie.

    Google Scholar 

  • Lucas, S. G. (1993a). The Shansiodon Biochron, nonmarine Middle Triassic of Pangaea. Albertiana, 11, 40–42.

    Google Scholar 

  • Lucas, S. G. (1993b). Vertebrate biochronology of the Triassic of China. In S. G. Lucas & M. Morales (Eds.), The nonmarine Triassic (pp. 301–306). New Mexico Museum of Natural History and Science Bulletin, 3.

    Google Scholar 

  • Lucas, S. G. (1996). Vertebrate biochronology of the Mesozoic of China. Memoirs of the Beijing Natural History Museum, 55, 110–148.

    Google Scholar 

  • Lucas, S. G. (1997). Dicynodon and Late Permian Pangaea. In W. Naiwen & J. Remane (Eds.), Proceedings of the 30th International Geological Congress (Vol. 11, pp. 133–141). Utrecht: VSP International Science Publishers.

    Google Scholar 

  • Lucas, S. G. (1998a). Toward a tetrapod biochronology of the Permian. In S. G. Lucas, J. W. Estep, & J. M. Hoffer (Eds.), Permian stratigraphy and paleontology of the Robledo Mountains, New Mexico (pp. 71–91). New Mexico Museum of Natural History and Science Bulletin, 12.

    Google Scholar 

  • Lucas, S. G. (1998b). Global Triassic tetrapod biostratigraphy and biochronology. Palaeogeography, Palaeoclimatology, Palaeoecology, 143, 347–384.

    Google Scholar 

  • Lucas, S. G. (1999). A tetrapod-based Triassic timescale. Albertiana, 22, 31–40.

    Google Scholar 

  • Lucas, S. G. (2001). Chinese fossil vertebrates. New York: Columbia University Press.

    Google Scholar 

  • Lucas, S. G. (2002). Tetrapods and the subdivision of Permian time. In L. V. Hillis, C. M. Henderson, & E. W. Bamber (Eds.), Carboniferous and Permian of the World (pp. 479–491). Canadian Society of Petroleum Geologists Memoir, 19.

    Google Scholar 

  • Lucas, S. G. (2005) Permian tetrapod faunachrons. In S. G. Lucas & K. E. Zeigler (Eds.), The nonmarine Permian (pp. 197–201). New Mexico Museum of Natural History and Science Bulletin, 30.

    Google Scholar 

  • Lucas, S. G. (2006). Global Permian tetrapod biostratigraphy and biochronology. In S. G. Lucas, G. Cassinis, & J. W. Schneider (Eds.), Non-marine Permian biostratigraphy and biochronology (Vol. 265, pp. 65–93). London: Geological Society, Special Publications.

    Google Scholar 

  • Lucas, S. G. (2009). Timing and magnitude of tetrapod extinctions across the Permo-Triassic bounary. Journal of Asian Earth Sciences, 36, 491–502.

    Google Scholar 

  • Lucas, S. G. (2010). The Triassic timescale based on nonmarine tetrapod biostratigraphy and biogeography. In S. G. Lucas (ed.), The Triassic timescale (Vol. 334, pp. 447–500). London: Geological Society, Special Publications.

    Google Scholar 

  • Lucas, S. G., & Wild, R. (1995). A Middle Triassic dicynodont from Germany and the biochronology of Triassic dicynodonts. Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie), 220, 1–16.

    Google Scholar 

  • Maisch., M. W. (1999). The tetrapods from the Late Permian of Tanzania in the collections of the Institut und Museum für Geologie und Paläontologie der Universität Tübingen, with special reference to the pristerodontian dicynodonts Rhachiocephalus and Pelanomodon. Unpublished Ph.D. thesis, Eberhard Karls Universität Tübingen, Tübingen.

    Google Scholar 

  • Maisch, M. W. (2000). Observations on Karoo vertebrates. Part 1. The taxonomic status of Rhachiocephalus usiliensis (von Huene, 1942) (Therapsida, Dicynodontia) from the Upper Permian Kawinga Formation of Tanzania. Neues Jahrbuch für Geologie und Paläontologie Monatshefte, 2000, 15–28.

    Google Scholar 

  • Maisch, M. W. (2001). Observations on Karoo and Gondwana vertebrates. Part 2: A new skull-reconstruction of Stahleckeria potens von Huene, 1935 (Dicynodontia, Middle Triassic) and a reconsideration of kannemeyeriiform phylogeny. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen, 220, 127–152.

    Google Scholar 

  • Maisch, M. W. (2002a). Observations on Karoo and Gondwana vertebrates. Part 4: The taxonomic status of the Late Permian rhachiocephalid Platycyclops crassus Broom, 1948 (Therapsida: Dicynodontia) from the South African Karoo. Neues Jahrbuch für Geologie und Paläontologie Monatshefte, 2002, 362–372.

    Google Scholar 

  • Maisch, M. W. (2002b). A new basal lystrosaurid dicynodont from the Upper Permian of South Africa. Palaeontology, 45, 343–359.

    Google Scholar 

  • Maisch, M. W. (2002c). Observation on Karoo and Gondwana vertebrates. Part 3: Notes on the gorgonopsians from the Upper Permian of Tanzania. Neues Jahrbuch für Geologie und Paläontologie Monatshefte, 2002, 237–251.

    Google Scholar 

  • Nicolas, M., & Rubidge, B. S. (2009). Assessing content and bias in South African Permo-Triassic Karoo tetrapod fossil collections. Palaeontologia Africana, 44, 13–20.

    Google Scholar 

  • Nicolas, M., & Rubidge, B. S. (2010). Changes in Permo-Triassic ecological representation in the Beaufort Group (Karoo Supergroup) of South Africa. Lethaia, 43, 45–59.

    Google Scholar 

  • Nopcsa, F. (1923). Die Familien der Reptilien. Berlin: Verlag von Gebrüder Borntraeger.

    Google Scholar 

  • Nyambe, I. A. (1999). Tectonic and climatic controls on sedimentation during deposition of the Sinakumbe Group and Karoo Supergroup, in the mid-Zambezi Valley Basin, southern Zambia. Journal of African Earth Sciences, 28, 443–463.

    Google Scholar 

  • Nyambe, I. A., & Utting, J. (1997). Stratigraphy and palynostratigraphy, Karoo Supergroup (Permian and Triassic), mid-Zambezi Valley, southern Zambia. Journal of African Earth Sciences, 24, 563–583.

    Google Scholar 

  • Ochev, V. G., & Shishkin, M. A. (1989). On the principles of global correlation of the continental Triassic on the tetrapods. Acta Palaeontologica Polonica, 34, 149–173.

    Google Scholar 

  • Owen, R. (1845). Report on the reptilian fossils of South Africa. Part I.—Description of certain fossil crania, discovered by A. G. Bain, Esq., in sandstone rocks at the south-eastern extremity of Africa, referable to different species of an extinct genus of Reptilia (Dicynodon), and indicative of a new tribe or sub-order of Sauria. Transactions of the Geological Society of London, Second Series, 7, 59–84.

    Google Scholar 

  • Owen, R. (1860a). On the orders of fossil and recent Reptilia and their distribution in time. Report of the British Association for the Advancement of Science, 1859, 153–166.

    Google Scholar 

  • Owen, R. (1860b). On some reptilian fossils from South Africa. Quarterly Journal of the Geological Society of London, 16, 49–63.

    Google Scholar 

  • Owen, R. (1876). Descriptive and illustrated catalogue of the fossil reptilia of South Africa in the collections of the British Museum. London: Taylor and Francis.

    Google Scholar 

  • Poole, D. F. G. (1956). The structure of the teeth of some mammal-like reptiles. Quarterly Journal of Microscopical Science, 97, 303–312.

    Google Scholar 

  • Ray, S. (1997). Some contributions to the lower Gondwanan stratigraphy of the Pranhita-Godavari Valley, Deccan, India. Journal of the Geological Society of India, 50, 633–640.

    Google Scholar 

  • Ray, S. (1999). Permian reptilian fauna from the Kundaram Formation, Pranhita-Godvari Valley, India. Journal of African Earth Sciences, 29, 211–218.

    Google Scholar 

  • Ray, S. (2000). Endothiodont dicynodonts from the Late Permian Kundaram Formation, India. Palaeontology, 43, 375–404.

    Google Scholar 

  • Ray, S. (2001). Small Permian dicynodonts from India. Paleontological Research, 5, 177–191.

    Google Scholar 

  • Ray, S. (2005). Lystrosaurus (Therapsida, Dicynodontia) from India: Taxonomy, relative growth and cranial dimorphism. Journal of Systematic Palaeontology, 3, 203–221.

    Google Scholar 

  • Ray, S., & Bandyopadhyay, S. (2003). Late Permian vertebrate community of the Pranhita-Godavari Valley, India. Journal of Asian Earth Sciences, 21, 643–654.

    Google Scholar 

  • Renaut, A. J. (2000). A re-evaluaiton of the cranial morphology and taxonomy of the Triassic dicynodont genus Kannemeyeria. Unpublished Ph.D. thesis, University of the Witwatersrand, Johannesburg.

    Google Scholar 

  • Renaut, A. J., & Hancox, P. J. (2001). Cranial description and taxonomic re-evaluation of Kannemeyeria argentinensis (Therapsida: Dicynodontia). Palaeontologia Africana, 37, 81–91.

    Google Scholar 

  • Renaut, A. J., Damiani, R. J., Yates, A. M., & Hancox, P. J. (2003). A taxonomic note concerning a dicynodont (Synapsida: Anomodontia) from the Middle Triassic of East Africa. Palaeontologia Africana, 39, 93–94.

    Google Scholar 

  • Roy Chowdhury, T. (1970). Two new dicynodonts from the Triassic Yerrapalli Formation of Central India. Palaeontology, 13, 132–144.

    Google Scholar 

  • Rubidge, B. S. (2005). Reuniting lost continents—fossil reptiles from the ancient Karoo and their wanderlust. South African Journal of Geology, 108, 135–172.

    Google Scholar 

  • Rubidge, B. S., & Sidor, C. A. (2001). Evolutionary patterns among Permo-Triassic therapsids. Annual Review of Ecology and Systematics, 32, 449–480.

    Google Scholar 

  • Rubidge, B. S., Johnson, M. R., Kitching, J. W., Smith, R. M. H., Keyser, A. W., & Groenewald, G. H. (1995). An introduction to the biozonation of the Beaufort Group. In B. S. Rubidge (Ed.), Biostratigraphy of the Beaufort Group (Karoo Supergroup) (pp. 1–2). South African Committee for Stratigraphy Biostratigraphic Series, 1.

    Google Scholar 

  • Seeley, H. G. (1889). Researches on the structure, organisation, and classification of the fossil Reptilia.—VI. On the anomodont Reptilia and their allies. Philosophical Transactions of the Royal Society of London B, 180, 215–296.

    Google Scholar 

  • Seeley, H. G. (1894). Researches on the structure, organisation, and classification of the fossil Reptilia.—Part IX., Section 1. On the Therosuchia. Philosophical Transactions of the Royal Society of London. Series B, Biological sciences, 185, 987–1018.

    Google Scholar 

  • Sidor, C. A., Angielczyk, K. D., Weide, D. M., Smith, R. M. H., Nesbitt, S. J., & Tsuji, L. A. (2010). Tetrapod fauna of the lowermost Usili Formation (Songea Group, Ruhuhu Basin) of southern Tanzania, with a new burnetiid record. Journal of Vertebrate Paleontology, 30, 696–703.

    Google Scholar 

  • Smith, R. M. H. (1987). Helical burrow casts of therapsid origin from the Beaufort Group (Permian) of South Africa. Palaeogeography, Palaeoclimatology, Palaeoecology, 60, 155–170.

    Google Scholar 

  • Smith, R. M. H. (1995). Changing fluvial environments across the Permian-Triassic boundary in the Karoo Basin, South Africa and possible causes of tetrapod extinctions. Palaeogeography, Palaeoclimatology, Palaeoecology, 117, 81–104.

    Google Scholar 

  • Smith, R. M. H., & Keyser, A. W. (1995). Biostratigraphy of the Cistecephalus Assemblage Zone. In B. S. Rubidge (Ed.), Biostratigraphy of the Beaufort Group (Karoo Supergroup) (pp. 23–28). South African Committee for Stratigraphy Biostratigraphic Series, 1.

    Google Scholar 

  • Smith, R. M. H., Eriksson, P. G., & Botha, W. J. (1993). A review of the stratigraphy and sedimentary environments of the Karoo-aged basins of Southern Africa. Journal of African Earth Sciences, 16, 143–169.

    Google Scholar 

  • Steyer, J. S. (2009). The geological and palaeontological exploration of Laos—following in the footsteps of J.B.H. Counillon. Journal of the Geological Society, London, 315, 25–32.

    Google Scholar 

  • Sues, H.-D., & Fraser, N. C. (2010). Triassic life on land. New York: Columbia University Press.

    Google Scholar 

  • Sullivan, C., & Reisz, R. R. (2005). Cranial anatomy and taxonomy of the Late Permian dicynodont Diictodon. Annals of Carnegie Museum, 74, 45–75.

    Google Scholar 

  • Sullivan, C., Reisz, R. R., & Smith, R. M. H. (2003). The Permian mammal-like herbivore Diictodon, the oldest known example of sexually dimorphic armament. Proceedings of the Royal Society of London B, 270, 173–178.

    Google Scholar 

  • Surkov, M. V. (2000). On the historical biogeography of Middle Triassic anomodonts. Paleontological Journal, 34, 84–88.

    Google Scholar 

  • Surkov, M. V., & Benton, M. J. (2004). The basicranium of dicynodonts (Synapsida) and its use in phylogenetic analysis. Palaeontology, 47, 619–638.

    Google Scholar 

  • Surkov, M. V., Kalandadze, N. N., & Benton, M. J. (2005). Lystrosaurus georgi, a dicynodont from the Lower Triassic of Russia. Journal of Vertebrate Paleontology, 25, 402–413.

    Google Scholar 

  • Tankard, A., Welsink, H., Aukes, P., Newton, R., & Stettler, E. (2009). Tectonic evolution of the Cape and Karoo basins of South Africa. Marine and Petroleum Geology, 26, 1379–1412.

    Google Scholar 

  • Toerien, M. J. (1954). Note on the systematic position of Compsodon, van H. Navorsinge van die Nasionale Museum, 1, 131–132.

    Google Scholar 

  • Tollman, S. M., Grine, F. E., & Hahn, B. D. (1980). Ontogeny and sexual dimorphism in Aulacephalodon (Reptilia, Anomodontia). Annals of the South African Museum, 81, 159–186.

    Google Scholar 

  • van Hoepen, E. C. N. (1934). Oor die indeling van die Dicynodontidae na aanleiding van nuew vorme. Paleontologiese Navorsing van die Nasionale Museum, 2, 67–101.

    Google Scholar 

  • Ward, P. D., Botha, J., Buick, R., De Kock, M. O., Erwin, D. H., Garrison, G. H., et al. (2005). Abrupt and gradual extinction among Late Permian land vertebrates in the Karoo Basin, South Africa. Science, 307, 709–714.

    Google Scholar 

  • Weide, D. M., Sidor, C. A., Angielczyk, K. D., & Smith, R. M. H. (2009). A new record of Procynosuchus delaharpeae (Therapsida: Cynodontia) from the Upper Permian Usili Formation, Tanzania. Palaeontologia Africana, 44, 21–26.

    Google Scholar 

  • Yeh, H.-K. (1959). New dicynodont from Sinokannemeyeria-fauna from Shansi. Vertebrata PalAsiatica, 3, 187–204.

    Google Scholar 

  • Yemane, K., & Kelts, K. (1990). A short review of palaeoenvironments for Lower Beaufort (Upper Permian) Karoo sequences from southern to central Africa: A major Gondwana lacustrine episode. Journal of African Earth Sciences, 10, 169–185.

    Google Scholar 

Download references

Acknowledgments

Fieldwork in Zambia in 2009 was made possible by a grant from the National Geographic Society (CRE 8571-08 to J.S.S.) and additional support from the Field Museum of Natural History. R.L.W.’s participation in the 2009 expedition was made possible by a grant from The Field Museum/IDP Foundation, Inc. African Training Fund (to K.D.A.). K. Mwamulowe and C. Chipote (NHCC) provided much helpful assistance in planning and carrying out the 2009 expedition, and in obtaining our temporary export permits. J.S.S. thanks the UMR 7207 of the CNRS (Paris) for additional support. A. Goulding and J. Menke made valuable contributions to the fieldwork and provided additional logistical support in Zambia. B. Rubidge and B. Zipfel (BP) arranged the loan of BP/1/3337, BP/1/3591, and BP/1/3603 to K.D.A. A. Shinya, L. Herzog, and C. Van Beek (Field Museum of Natural History) prepared many of the specimens figured in this paper, including all of the figured material from the 2009 expedition. We thank the anonymous reviewers for their helpful comments.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Kenneth D. Angielczyk .

Editor information

Editors and Affiliations

Rights and permissions

Reprints and permissions

Copyright information

© 2014 Springer Science+Business Media Dordrecht

About this chapter

Cite this chapter

Angielczyk, K.D., Steyer, JS., Sidor, C.A., Smith, R.M.H., Whatley, R.L., Tolan, S. (2014). Permian and Triassic Dicynodont (Therapsida: Anomodontia) Faunas of the Luangwa Basin, Zambia: Taxonomic Update and Implications for Dicynodont Biogeography and Biostratigraphy. In: Kammerer, C., Angielczyk, K., Fröbisch, J. (eds) Early Evolutionary History of the Synapsida. Vertebrate Paleobiology and Paleoanthropology. Springer, Dordrecht. https://doi.org/10.1007/978-94-007-6841-3_7

Download citation

Publish with us

Policies and ethics