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A Non-Mammaliaform Cynodont from the Upper Triassic of South Africa: a Therapsid Lazarus Taxon?

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A Non-Mammaliaform Cynodont from the Upper Triassic of South Africa: a Therapsid Lazarus Taxon? PALAEONTOLOGIA AFRICANA Volume 42 May 2007 Annals of the Bernard Price Institute for Palaeontological Research VOLUME 42, 2007 AFRICANA PALAEONTOLOGIA Supported by PALAEONTOLOGICAL SCIENTIFIC TRUST PALAEONTOLOGICAL SCIENTIFIC TRUST ISSN 0078-8554 SCHOOL OF GEOSCIENCES BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH Academic Staff Senior Administrative Secretary Director and Chair of Palaeontology S.C. Tshavhumbwe B.S. Rubidge BSc (Hons), MSc (Stell), PhD (UPE) Assistant Research Technician Deputy Director C.B. Dube M.K. Bamford BSc (Hons), MSc, PhD (Witwatersrand) Technician/Fossil Preparator Research Officers S. Jirah A.M. Yates, BSc (Adelaide), BSc (Hons), PhD (La Trobe) P.R. Mukanela Collections Curator G. Ndlovu B. Zipfel NHD Pod., NHD PS Ed. (TWR), BSc (Hons) T. Nemavhundi (Brighton), PhD (Witwatersrand) J.N. Sithole Post Doctoral Fellows S. Tshabalala F. Abdala BSc, PhD (UNT, Argentina) R. Govender BSc (Hons) UP, MSc, PhD (Witwatersrand) Custodian, Makapansgat Sites J. Maluleke Editorial Panel M.K. Bamford: Editor Honorary Staff L.R. Backwell: Associate Editor Honorary Professor of Palaeoanthropology B.S. Rubidge: Associate Editor P.V. Tobias OSc, OMSG (S.Afr.), FRS, FRCP, MBBCh, PhD., A.M. Yates: Associate Editor DSc (Witwatersrand), Hon. ScD (Cantab, Pennsylvania), Hon. DSc (Natal, U West., Ont., Alberta, Cape Town, Consulting Editors Guelph, UNISA, Durban-Westville, Pennsylvania, Wits, Dr J.A. Clack (Museum of Zoology, University of Mus. d’Hist Naturelle – Paris, Barcelona, Turin, Charles Cambridge, Cambridge, U.K.) U, Prague, Stellenbosch, Unitra, Fribourg), For. Assoc. Dr H.C. Klinger (South African Museum, Cape Town) NAS, Hon. FRSSA, Hon. FCMSA, FASSA Dr K. Padian (University of California, Berkeley, California, U.S.A.) Honorary Research Associates Dr K.M. Pigg (Arizona State University, Arizona, U.S.A.) C.K. Brain BSc, PhD (UCT), DSc (Witwatersrand), Hon. DSc (UCT, Natal, Pret., Witwatersrand), FZS, FRSSA Prof. L. Scott (University of the Free State, Bloemfontein) F.E. Grine BA (Hons) (Washington & Jefferson College), Dr R.M.H. Smith (South African Museum, Cape Town) PhD (Witwatersrand) Dr J.F. Thackeray (Northern Flagship Institution, Pretoria) J.M. Maguire BSc (Natal), BSc (Hons), PhD Technical and Support Staff (Witwatersrand) Principal Technician I.R. McLachlan BSc (Hons) (Witwatersrand) R. McRae-Samuel C.A. Sidor BSc (Trinity College) MSc, PhD (Univ. Chicago) INSTITUTE FOR HUMAN EVOLUTION Academic Staff Honorary Research Associates S.E. Churchill PhD (New Mexico) Reader A. Keyser BSc (Pret), MSc (Pret), PhD (Witwatersrand) L.R. Berger BA (Hons) (GA Southern), PhD R.S. Kidd Dip. Pod. Med. (Salford), BA (Hons) (Open (Witwatersrand) University), PhD (Western Australia) P. Schmid PhD (Zürich) Research Officer P.S. Ungar BA (SUNY Binghamton), MA, PhD (SUNY, L.R. Backwell BA (Hons), MSc , PhD (Witwatersrand) Stony Brook) PALAEONTOLOGIA AFRICANA ANNALS OF THE BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH UNIVERSITY OF THE WITWATERSRAND ISSN 0078-8554 Volume 42, May 2007 CONTENTS Articles Fourie, H. & Rubidge, B.S. — The postcranial skeletal anatomy of the 1 therocephalian Regisaurus (Therapsida: Regisauridae) and its utilization for biostratigraphic correlation Abdala, F., Damiani, R., Yates, A. & Neveling, J. — A non-mammaliaform 17 cynodont from the Upper Triassic of South Africa: a therapsid Lazarus taxon? Klinger, H.C. & McMillan, I.K. — Ammonites from offshore deposits near 25 Bogenfels, Namibia Pickford, M. — New mammutid proboscidean teeth from the Middle Miocene 29 of tropical and southern Africa Lewis, P.J. & Simons, E.L. — Morphological trends in the molars of fossil 37 rodents from the Fayum Depression, Egypt Schubert, B.W. — Dental mesowear and the palaeodiets of bovids from 43 Makapansgat Limeworks Cave, South Africa Eisenmann, V.& Geraads D., — Hipparion pomeli sp. nov from the late Pliocene 51 of Ahl al Oughlam, Morocco, and a revision of the relationships of Pliocene and Pleistocene African hipparions Hartstone-Rose, A., De Ruiter, D.J., Berger, L.R. & Churchill, S.E. —A 99 sabre-tooth felid from Coopers Cave (Gauteng, South Africa) and its implications for Megantereon (Felidae: Machairodontinae) taxonomy Broadley, D.G. — Description of an anomalous tortoise (Reptilia: Testudinidae) 109 from the Early Holocene of Zimbabwe Technical Report McCrae, C. & Potze, S. — A fresh look at chemical fossil extraction 115 Abstracts 15th Biennial Meeting of the Palaeontological Society of Southern Africa, 117 Albany Museum and Rhodes University, 7–10 September 2006 © 2007 BERNARD PRICE INSTITUTE for PALAEONTOLOGICAL RESEARCH School of Geosciences University of the Witwatersrand Johannesburg ACKNOWLEDGEMENTS The Bernard Price Institute for Palaeontological Research gratefully acknowledges financial support for its programmes by THE COUNCIL’S RESEARCH COMMITTEE, UNIVERSITY OF THE WITWATERSRAND NATIONAL RESEARCH FOUNDATION (NRF) DEPARTMENT OF SCIENCE AND TECHNOLOGY (DST) and the PALAEONTOLOGICAL SCIENTIFIC TRUST (PAST) for publication of this journal Pre-press production by Isteg Scientific Publications, Irene Printed in South Africa by Ultra Litho (Pty) Ltd, Heriotdale, Johannesburg The postcranial skeletal anatomy of the therocephalian Regisaurus (Therapsida: Regisauridae) and its utilization for biostratigraphic correlation Heidi Fourie* & Bruce S. Rubidge Bernard Price Institute for Palaeontological Research, University of the Witwatersrand, Johannesburg, Private Bag 3, WITS, 2050 South Africa Received 25 August 2006. Accepted 18 April 2007 The postcranial morphology of the therocephalian genus Regisaurus from the Lystrosaurus Assemblage Zone of South Africa is described. The remarkably complete state of preservation of the vertebral column has, for the first time, provided a full vertebral count for a therocephalian and demonstrates that it is possible to differentiate between cervical, thoracic, lumbar, sacral and caudal vertebrae. It is demonstrated that some postcranial elements can be used to identify particular therocephalian groups and will be of use in biostratigraphic studies in areas where cranial remains have not been found. A slender scapula, low scapular ridge, shallow scapular depression, short and broad interclavicle, oval sternum, and a small obturator foramen are characteristics of therocephalians known from the Cistecephalus, Dicynodon and Lystrosaurus assemblage zones of the Beaufort Group of South Africa. Keywords: Therapsida, Therocephalia, Regisaurus, postcranium. INTRODUCTION Bethulie, South Africa. Proterosuchus and Lystrosaurus are Therocephalians form a significant component of the co-occurring genera from the same horizon at this locality, therapsid faunas from the Karoo of South Africa, and whereas Dicynodon (=Daptocephalus) and Moschorhinus together with dicynodonts are the therapsid clade that are recorded from stratigraphically lower levels (Kitching have the longest stratigraphic range in the rocks of the 1977). Karoo Supergroup. They also have a wide geographic The specimen was preserved in a single calcareous range being known from Permian and Triassic rocks of nodule and was exposed by mechanical preparation using southern and eastern Africa (Boonstra 1935, 1969; compressed air-driven engravers fitted with tungsten Drysdall & Kitching 1963; Kitching 1977; Keyser & Brink carbide tips. Because of the uniquely complete nature of 1979), China (Li & Cheng 1995), Russia (Tatarinov 1974), the specimen great care was taken to preserve the original and Antarctica (Colbert & Kitching1981). Many thero- bone surface. For comparative purposes several thero- cephalian taxa from South Africa have been described, cephalian specimens that have postcranial elements and although several relatively recent descriptions on preserved were studied in various museum collections the cranial morphology are known (e.g. van den Heever (Table 1). 1987; 1994; Durand 1991; Hillenius 1992, 1994), studies on the postcranium of this important group have POSTCRANIAL DESCRIPTION lagged behind. This paper describes a unique and al- The skeleton of BP/1/3973 is almost complete, lacking most completely preserved therocephalian skeleton only the right manus, left pes, left femur, left ilium and (BP/1/3973). The presence of a long and high lacrimal; portion of the left tibia and right tibia and fibula. Parts of jugal meets the postorbital posteriorly and makes up the the pelvic girdle were disarticulated prior to fossilization ventral portion of the postorbital bar; parietal present on and are preserved on the left side of the body close to the dorsal part of occipital surface; upper dental formula of distal tibia and fibula (Figs 1B, 2B). As the holotype I6 C1 PC10; absence of precanines; palatal teeth on preserves only parts of the ilium and hind limb and pterygoid boss; and apparent absence of a pineal foramen elements of the shoulder girdle, specimen BP/1/3973 has enables us to identify the specimen as Regisaurus much to contribute to our understanding of therocephalian (Mendrez 1975). Mendrez (1972) described the holotype postcranial anatomy. In the following description the ‘in of Regisaurus (BP/1/5394) on cranial morphology alone life’ dorsal, ventral, lateral, and medial orientation of the even though isolated elements of the hind limb, pelvic different bones is assumed. and pectoral girdles, and vertebral fragments are preserved. Kemp (1978) described the ilium and hind limb of the Vertebrae (Fig. 3) same specimen and made conclusions on the stance and A remarkable feature of BP/1/3973 is
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