A reevaluation of Brachyprosopus broomi and Chelydontops altidentalis, dicynodonts (Therapsida, Anomodontia) from the middle Permian Tapinocephalus Assemblage Zone of the Karoo Basin, South Africa

LITERATURE CITED

• Angielczyk, K. D. 2001. Preliminary phylogenetic analysis and stratigraphic congruence of the dicynodont anomodonts (Synapsida: Therapsida). Palaeontologia africana 37:53–79.
   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.
   Web of Science ®Google Scholar
• Angielczyk, K. D. 2004. Phylogenetic evidence for and implications of a dual origin of propaliny in anomodont therapsids (Synapsida). Paleobiology 30:268–296.
   Web of Science ®Google Scholar
• Angielczyk, K. D. 2007. New specimens of the Tanzanian dicynodont “Cryptocynodon” parringtoni von Huene, 1942 (Therapsida, Anomodontia), with an expanded analysis of Permian dicynodont phylogeny. Journal of Vertebrate Paleontology 27:116–131.
   Web of Science ®Google Scholar
• Angielczyk, K. D., and C. B. Cox. 2015. Distinctive emydopoid dicynodont (Therapsida, Anomodontia) mandibles from the Permian Ruhuhu and Usili formations (Songea Group), Ruhuhu Basin, Tanzania. Journal of Vertebrate Paleontology 35:6, e1008699, DOI: 10.1080/02724634.2015.1008699.
   Web of Science ®Google Scholar
• Angielczyk, K. D., and A. A. Kurkin. 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.
   Web of Science ®Google Scholar
• Angielczyk, K. D., and B. S. Rubidge. 2009. The Permian dicynodont Colobodectes cluveri (Therapsida, Anomodontia), with notes on its ontogeny and stratigraphic range in the Karoo Basin, South Africa. Journal of Vertebrate Paleontology 29:1162–1173.
   Web of Science ®Google Scholar
• Angielczyk, K. D., and B. S. Rubidge. 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.
   Web of Science ®Google Scholar
• Angielczyk, K. D., and B. S. Rubidge. 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.
   Web of Science ®Google Scholar
• Angielczyk, K. D., and C. S. Sullivan. 2008. Diictodon feliceps (Owen, 1876), a dicynodont (Therapsida, Anomodontia) species with a Pangaean distribution. Journal of Vertebrate Paleontology 28:788–802.
   Web of Science ®Google Scholar
• Angielczyk, K. D., and M. L. Walsh. 2008. Patterns in the evolution of nares size and secondary palate length in anomodont therapsids (Synapsida): implication for hypoxia as a cause of the end-Permian tetrapod extinctions. Journal of Paleontology 82:528–542.
   Web of Science ®Google Scholar
• Angielczyk, K. D., J. Fröbisch, and R. M. H. Smith. 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., C. A. Sidor, S. J. Nesbitt, R. M. H. Smith, and L. A. Tsuji. 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.
   Web of Science ®Google Scholar
• Angielczyk, K. D., S. Huertas, R. M. H. Smith, N. J. Tabor, C. A. Sidor, J. S. Steyer, and L. A. Tsuji. A. 2014. New dicynodonts (Therapsida, Anomodontia) and updated tetrapod stratigraphy of the Permian Ruhuhu Formation (Songea Group, Ruhuhu Basin) of southern Tanzania. Journal of Vertebrate Paleontology 34:1408–1426.
   Web of Science ®Google Scholar
• Barry, T. H. 1967. The cranial morphology of the Permo-Triassic anomodont Pristerodon buffaloensis with special reference to the neural endocranium and the visceral arch skeleton. Annals of the South African Museum 50:131–161.
   Google Scholar
• Barry, T. H. 1974. A new dicynodont ancestor from the upper Ecca (lower middle Permian) of South Africa. Annals of the South African Museum 64:117–136.
   Google Scholar
• Barry, T. H. 1975. Affinities and systematic position of the South African lower middle Permian dicynodonts (Therapsida: Dicynodontidae); pp. 475–479 in K. S. W. Campbell (ed.), Gondwana Geology: Papers Presented at the Third Gondwana Symposium, Canberra, Australia. Australian National University Press, Canberra, Australia.
   Google Scholar
• Boonstra, L. D. 1948. On the anomodont reptiles from the Tapinocephalus-Zone of the Karroo System; pp. 57–64 in A. L. Du Toit (ed.), Robert Broom Commemorative Volume. Royal Society of South Africa, Cape Town, South Africa.
   Google Scholar
• Boonstra, L. D. 1963. Early dichotomies in the therapsids. South African Journal of Science 59:176–195.
   Google Scholar
• Boonstra, L. D. 1972. Discard the names Theriodontia and Anomodontia: a new classification of Therapsida. Annals of the South African Museum 59:315–338.
   Google Scholar
• Boos, A. D. S., C. L. Schultz, C. S. Vega, and J. J. Aumond. 2013. On the presence of the late Permian dicynodont Endothiodon in Brazil. Palaeontology 56:837–848.
   Web of Science ®Google Scholar
• Botha, J., and K. D. Angielczyk. 2007. An integrative approach to distinguishing the late Permian dicynodont species Oudenodon bainii and Tropidostoma microtrema (Therapsida: Anomodontia). Palaeontology 50:1175–1209.
   Web of Science ®Google Scholar
• Bremer, K. 1988. The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution 42:795–803.
   PubMed Web of Science ®Google Scholar
• Bremer, K. 1994. Branch support and tree stability. Cladistics 10:295–304.
   Web of Science ®Google Scholar
• Brink, A. S. 1983. Chelydontops altidentalis Cluver, 1975. Illustrated bibliographic catalogue of the Synapsida. Geological Survey of South Africa Handbook 10:J212A221A.
   Google Scholar
• Brink, A. S., and A. W. Keyser. 1983. Pristerodon mackayi Huxley, 1868. Illustrated bibliographic catalogue of the Synapsida. Geological Survey of South Africa Handbook 10:J212A231A.
   Google Scholar
• Broili, F., and J. Schröder. 1935. Beobachtungen an Wirbeltieren der Karrooformation. XII. Über einige primitive Anomodontier-Schädel aus den unteren Beaufort-Schichten. Sitzungsberichte der Bayerischen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Abteilung 1935:223–278.
   Google Scholar
• Broom, R. 1905a. On the use of the term Anomodontia. Albany Museum Records 1:266–269.
   Google Scholar
• Broom, R. 1905b. On the structure and affinities of the endothiodont reptiles. Transactions of the South African Philosophical Society 15:259–282.
   Google Scholar
• Broom, R. 1932. The Mammal-like Reptiles of South Africa and the Origin of Mammals. H. F. and G. Witherby, London, 376 pp.
   Google Scholar
• Castanhinha, R., R. Araújo, L. C. Júnior, K. D. Angielczyk, G. G. Martins, R. M. S. Martins, C. Chaouiya, F. Backmann, and F. Wilde. 2013. Bringing dicynodonts back to life: paleobiology and anatomy of a new emydopoid genus from the upper Permian of Mozambique. PLoS ONE 8:e80974.
   PubMed Web of Science ®Google Scholar
• Cluver, M. A. 1970. The palate and mandible in some specimens of Dicynodon testudirostris Broom and Haughton (Reptilia, Therapsida). Annals of the South African Museum 56:133–153.
   Google Scholar
• Cluver, M. A. 1971. The cranial morphology of the dicynodont Lystrosaurus. Annals of the South African Museum 56:155–274.
   Google Scholar
• Cluver, M. A. 1975. A new dicynodont reptile from the Tapinocephalus Zone (Karoo System, Beaufort Series) of South Africa, with evidence of the jaw adductor musculature. Annals of the South African Museum 67:7–23.
   Google Scholar
• Cluver, M. A., and G. M. King. 1983. A reassessment of the relationships of Permian Dicynodontia (Reptilia, Therapsida) and a new classification of dicynodonts. Annals of the South African Museum 91:191–273.
   Google Scholar
• Cole, D. I., and P. E. Wipplinger. 2001. Sedimentology and molybdenum potential of the Beaufort Group in the main Karoo Basin, South Africa. Memoirs of the Council for Geoscience 80:1–225.
   Google Scholar
• Cox, C. B. 1959. On the anatomy of a new dicynodont genus with evidence of the position of the tympanum. Proceedings of the Zoological Society of London 132:321–367.
   Google Scholar
• Cox, C. B. 1962. A natural cast of the inner ear of a dicynodont. American Museum Novitates 2116:1–6.
   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.
   Web of Science ®Google Scholar
• Cox, C. B., and K. D. Angielczyk. 2015. A new endothiodont dicynodont (Therapsida, Anomodontia) from the Permian Ruhuhu Formation (Songea Group) of Tanzania and its feeding system. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2014.935388.
   Web of Science ®Google Scholar
• Crompton, A. W., and N. Hotton III. 1967. Functional morphology of the masticatory apparatus of two dicynodonts (Reptilia, Therapsida). Postilla 109:1–51.
   Google Scholar
• Day, M. O. 2013. Middle Permian continental biodiveristy changes as reflected in the Beaufort Group of South Africa: a bio- and lithostratigraphic review of the Eodicynodon, Tapinocephalus and Pristerognathus Assemblage Zones. Unpublished Ph.D. dissertation, University of the Witwatersrand, Johannesburg, South Africa, 387 pp.
   Google Scholar
• Day, M. O., and B. S. Rubidge. 2014. A brief lithostratigraphic review of the Abrahamskraal and Koonap formations of the Beaufort Group, South Africa: towards a basin-wide stratigraphic scheme for the middle Permian Karoo. Journal of African Earth Sciences 100:227–242.
   Web of Science ®Google Scholar
• Day, M., B. Rubidge, J. Almond, and S. Jirah. 2013. Biostratigraphic correlation in the Karoo: the case of the middle Permian parareptile Eunotosaurus. South African Journal of Science 109:68–71.
   Web of Science ®Google Scholar
• Day, M. O., J. Ramezani, S. A. Bowring, P. M. Sadler, D. H. Erwin, F. Abdala, and B. S. Rubidge. 2015a. When and how did the terrestrial mid-Permian mass extinction occur? Evidence from the tetrapod record of the Karoo Basin, South Africa. Proceedings of the Royal Society Series B 282:20150834.
   Google Scholar
• Day, M. O., S. Güven, F. Abdala, S. Jirah, B. Rubidge, and J. Almond. 2015b. Youngest dinocephalian fossils extend the Tapinocephalus zone, Karoo Basin, South Africa. South African Journal of Science 111:78–82.
   Web of Science ®Google Scholar
• Ewer, R. F. 1961. The anatomy of the anomodont Daptocephalus leoniceps (Owen). Proceedings of the Zoological Society of London 136:375–402.
   Google Scholar
• Fourie, H. 1993. A detailed description of the internal structure of the skull of Emydops (Therapsida: Dicynodontia). Palaeontologia africana 30:103–111.
   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 150:117–114.
   Web of Science ®Google Scholar
• Fröbisch, J., and R. R. Reisz. 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.
   Web of Science ®Google Scholar
• Fröbisch, J., K. D. Angielczyk, and C. A. Sidor. 2010. The Triassic dicynodont Kombuisia (Synapsida, Anomodontia) from Antarctica, a refuge from the terrestrial Permian-Triassic extinction. Naturwissenschaften 97:187–196.
   PubMed Web of Science ®Google Scholar
• Goloboff, P. A., J. S. Farris, and K. C. Nixon. 2008. TNT, a free program for phylogenetic analysis. Cladistics 24:774–786.
   Web of Science ®Google Scholar
• Goloboff, P. A., C. I. Mattoni, and A. S. Quinteros. 2006. Continuous characters analyzed as such. Cladistics 22:589–601.
   PubMed Web of Science ®Google Scholar
• Goloboff, P. A., J. S. Farris, M. Källersjö, B. Oxelman, M. J. Ramírez, and C. A. Szumik. 2003. Improvements to resampling measures of group support. Cladistics 19:324–332.
   Web of Science ®Google Scholar
• Jirah, S., and B. S. Rubidge. 2014. Refined stratigraphy of the middle Permian Abrahamskraal Formation (Beaufort Group) in the southern Karoo Basin. Journal of African Earth Sciences 100:121–135.
   Web of Science ®Google Scholar
• Kammerer, C. F., K. D. Angielczyk, and J. Fröbisch. 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.
   Web of Science ®Google Scholar
• Kammerer, C. F., K. D. Angielczyk, and J. Fröbisch. 2015. Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the late Permian of South Africa. Fossil Record 18:43–55.
   Web of Science ®Google Scholar
• Kammerer, C. F., J. Fröbisch, and K. D. Angielczyk. 2013. On the validity and phylogenetic position of Eubrachiosaurus browni, a kannemeyeriiform dicynodont (Anomodontia) from Triassic North America. PloS ONE 8:e64203.
   PubMed Web of Science ®Google Scholar
• Kemp, T. S. 1969. The atlas-axis complex of the mammal-like reptiles. Journal of Zoology 159:223–248.
   Web of Science ®Google Scholar
• Keyser, A. W. 1973. 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. 1974. Evolutionary trends in Triassic Dicynodontia. Palaeontologia africana 17:57–68.
   Google Scholar
• Keyser, A. W. 1993. A re-evaluation of the smaller Endothiodontidae. Geological Survey of South Africa Memoir 82:1–53.
   Google Scholar
• King, G. M. 1981. The functional anatomy of a Permian dicynodont. Philosophical Transactions of the Royal Society Series B 291:243–322.
   Web of Science ®Google Scholar
• King, G. M. 1988. Anomodontia; in Handbuch der Paläoherpetologie, 17 C. Gustav Fischer Verlag, Stuttgart, Germany, 174 pp.
   Google Scholar
• King, G. M. 1990. The Dicynodonts: A Study in Palaeobiology. Chapman and Hall, London, 233 pp.
   Google Scholar
• King, G. M. 1994. The early anomodont Venjukovia and the evolution of the anomodont skull. Journal of Zoology 232:651–673.
   Web of Science ®Google Scholar
• King, G. M., and B. S. Rubidge. 1993. A taxonomic revision of small dicynodonts with postcanine teeth. Zoological Journal of the Linnean Society 107:131–154.
   Web of Science ®Google Scholar
• King, G. M., B. W. Oelofsen, and B. S. Rubidge. 1989. The evolution of the dicynodont feeding system. Zoological Journal of the Linnean Society 96:185–211.
   Web of Science ®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
• Latimer, E. M., C. E. Gow, and B. S. Rubidge. 1995. Dentition and feeding niche of Endothiodon (Synapsida, Anomodontia). Palaeontologia africana 32:75–82.
   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. Ph.D. dissertation, Eberhard Karls Universität Tübingen, Tübingen, Germany, 350 pp.
   Google Scholar
• Modesto, S. P., B. S. Rubidge, and J. Welmann. 2002. A new dicynodont therapsid from the lowermost Beaufort Group, upper Permian of South Africa. Canadian Journal of Earth Sciences 39:1755–1765.
   Web of Science ®Google Scholar
• Modesto, S. P., B. Rubidge, I. Visser, and J. Welman. 2003. A new basal dicynodont from the upper Permian of South Africa. Palaeontology 46:211–223.
   Web of Science ®Google Scholar
• Nehls, M., C. Kammerer, and J. Fröbisch. 2014. CT reconstruction and systematic position of the enigmatic dicynodont Prodicynodon beaufortensis. Journal of Vertebrate Paleontology 34(3, Supplement):194–195.
   Google Scholar
• Nicolas, M. V. 2007. Tetrapod biodiversity through the Permo-Triassic Beaufort Group (Karoo Supergroup) of South Africa. Unpublished Ph.D. dissertation, University of the Witwatersrand, Johannesburg, South Africa, 312 pp.
   Google Scholar
• Nopsca, F. 1923. Die Familien der Reptilien. Verlag von Gebrüder Borntraeger, Berlin, Germany, 210 pp.
   Google Scholar
• Olson, E. C. 1937. The skull structure of a new anomodont. Journal of Geology 45:851–858.
   Google Scholar
• Olson, E. C. 1944. Origin of mammals based upon cranial morphology of the therapsid suborders. Geological Society of America Special Papers 55:1–136.
   Google Scholar
• Owen, R. 1860. On the orders of fossil and recent Reptilia, and their distribution in time. Report of the Twenty-Ninth Meeting of the British Association for the Advancement of Science, 1859:153–166.
   Google Scholar
• Ray, S. 2000. Endothiodont dicynodonts from the late Permian Kundaram Formation, India. Palaeontology 43:375–405.
   Web of Science ®Google Scholar
• Rowe, T. 1980. The morphology, affinities, and age of the dicynodont reptile Geikia elginensis; pp. 269–294 in L. L. Jacobs (ed.), Aspects of Vertebrate History. Museum of Northern Arizona Press, Flagstaff, Arizona.
   Google Scholar
• Rubidge, B. S. 1984. The cranial morphology and palaeoenvironment of Eodicynodon Barry (Therapsida: Dicynodontia). Navorsinge van die Nasionale Museum Bloemfontein 4:326–402.
   Google Scholar
• Rubidge, B. S. 1988. A palaeontological and palaeoenvironmental synthesis of the Permian Ecca—Beaufort contact in the southern Karoo between Prince Albert and Rietbron, Cape Province, South Africa. Unpublished Ph.D. dissertation, University of Port Elizabeth, Port Elizabeth, South Africa, 349 pp.
   Google Scholar
• Rubidge, B. S. 1990a. Redescription of the cranial morphology of Eodicynodon oosthuizeni (Therapsida: Dicynodontia). Navorsinge van die Nasionale Museum Bloemfontein 7:1–25.
   Google Scholar
• Rubidge, B. S. 1990b. The cranial morphology of a new species of the genus Eodicynodon (Therapsida, Dicynodontia). Navorsinge van die Nasionale Museum Bloemfontein 7:29–41.
   Google Scholar
• Rubidge, B. S. (ed.). 1995. Biostratigraphy of the Beaufort Group (Karoo Supergroup). South African Committee for Stratigraphy Biostratigraphic Series 1:1–46.
   Google Scholar
• Rubidge, B., and K. Angielczyk. 2009. Stratigraphic ranges of Tapinocephalus Assemblage Zone dicynodonts: implications for middle Permian continental biostratigraphy. Palaeontologia africana 44:134–135.
   Google Scholar
• Rubidge, B. S., G. M. King, and P. J. Hancox. 1994. The postcranial skeleton of the earliest dicynodont synapsid Eodicynodon from the upper Permian of South Africa. Palaeontology 37:397–408.
   Web of Science ®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 185:987–1018.
   Google Scholar
• Simon, R. V., C. A. Sidor, K. D. Angielczyk, and R. M. H. Smith. 2010. First record of a tapinocephalid (Therapsida: Dinocephalia) from the Ruhuhu formation (Songea Group) of southern Tanzania. Journal of Vertebrate Paleontology 30:1289–1293.
   Web of Science ®Google Scholar
• Smith, R., B. Rubidge, and M. van der Walt. 2012. Therapsid biodiversity patterns and paleoenvironments of the Karoo Basin, South Africa; pp. 30–62 in A. Chinsamy-Turan (ed.), Forerunners of Mammals: Radiation, Histology, Biology. Indiana University Press, Bloomington, Indiana.
   Google Scholar
• Sollas, I. B. J., and W. J. Sollas. 1914. A study of the skull of a Dicynodon by means of serial sectioning. Philosophical Transactions of the Royal Society Series B 204:201–225.
   Google Scholar
• Strong, E. E., and D. Lipscomb. 2000. Character coding and inapplicable data. Cladistics 15:363–371.
   Web of Science ®Google Scholar
• Sullivan, C., and R. R. Reisz. 2005. Cranial anatomy and taxonomy of the late Permian dicynodont Diictodon. Annals of Carnegie Museum 74:45–75.
   Web of Science ®Google Scholar
• Sullivan, C., R. R. Reisz, and R. M. H. Smith. 2003. The Permian mammal-like herbivore Diictodon, the oldest known example of sexually dimorphic armament. Proceedings of the Royal Society Series B 270:173–178.
   Web of Science ®Google Scholar
• Toerien, M. J. 1953. The evolution of the palate in South African Anomodontia and its classificatory significance. Palaeontologia africana 1:49–117.
   Google Scholar
• Van der Walt, M., M. Day, B. Rubidge, A. K. Cooper, and I. Netterberg. 2010. A new GIS-based biozone map of the Beaufort Group (Karoo Supergroup), South Africa. Palaeontologia africana 45:1–5.
   Google Scholar
• Van Hoepen, E. C. N. 1934. Oor die indeling van die Dicynodontidae na aanleidung van nuew vorme. Palaeontologiese Navorsing van die Nasionale Museum 2:67–101.
   Google Scholar
• Viglietti, P. A., R. M. H. Smith, K. D. Angielczyk, C. F. Kammerer, J. Fröbisch, and B. S. Rubidge. 2016. The Daptocephalus Assemblage Zone (Lopingian), South Africa: A proposed biostratigraphy based on a new compilation of stratigraphic ranges. Journal of African Earth Sciences :153–164.
   Google Scholar
• Handling editor: J. Botha-Brink.
   Google Scholar