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PALAEONTOLOGIA AFRICANA Volume 44 December 2009 Annals of the Bernard Price Institute for Palaeontological Research

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PALAEONTOLOGIA AFRICANA Volume 44 December 2009 Annals of the Bernard Price Institute for Palaeontological Research PALAEONTOLOGIA AFRICANA Volume 44 December 2009 Annals of the Bernard Price Institute for Palaeontological Research VOLUME 44, 2009 AFRICANA PALAEONTOLOGIA Supported by PALAEONTOLOGICAL SCIENTIFIC TRUST PALAEONTOLOGICAL SCIENTIFIC TRUST ISSN 0078-8554 SCHOOL OF GEOSCIENCES BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH Academic Staff Dr K. Padian (University of California, Berkeley, Director and Chair of Palaeontology California, U.S.A.) B.S. Rubidge BSc (Hons), MSc (Stell), PhD (UPE) Dr K.M. Pigg (Arizona State University, Arizona, U.S.A.) Deputy Director Prof. L. Scott (University of the Free State, Bloemfontein) M.K. Bamford BSc (Hons), MSc, PhD (Witwatersrand) Dr R.M.H. Smith (South African Museum, Cape Town) Senior Research Officers Technical and Support Staff F. Abdala BSc, PhD (UNT, Argentina) Principal Technician A.M. Yates, BSc (Adelaide), BSc (Hons), PhD (La Trobe) R. McRae-Samuel Research Officer Senior Administrative Secretary L.R. Backwell BA (Hons), MSc, PhD (Witwatersrand) S.C. Tshishonga Collections Curator Assistant Research Technician B. Zipfel NHD Pod., NHD PS Ed. (TWR), BSc (Hons) C.B. Dube (Brighton), PhD (Witwatersrand) Technician/Fossil Preparator Post Doctoral Fellows P. Chakane R. Mutter BSc, MSc, PhD (Zurich, Switzerland) S. Jirah F. Neumann BSc, MSc, PhD (Friedrich-Wilhelms-Univer- P.R. Mukanela sity, Bonn) G. Ndlovu D. Steart BSc, DipEd (La Trobe University), PhD (Victoria T. Nemavhundi University of Technology, Australia) S. Tshabalala Editorial Panel Custodian, Makapansgat Sites M.K. Bamford: Editor S. Maluleke L.R. Backwell: Associate Editor Honorary Staff B.S. Rubidge: Associate Editor Honorary Research Associates A.M. Yates: Associate Editor K. Angielczyk BSc (Univ of Michigan, Ann Arbor), PhD Consulting Editors (Univ California, Berkeley) Dr J.A. Clack (Museum of Zoology, University of P.J. Hancox BSc(Hons), MSc, PhD (Witwatersrand) Cambridge, Cambridge, U.K.) R. Reisz BSc, MSc, PhD (McGill Univ., Montreal) Dr H.C. Klinger (South African Museum, Cape Town) C.A. Sidor BSc (Trinity College) MSc, PhD (Univ. Chicago) INSTITUTE FOR HUMAN EVOLUTION Academic Staff T. Makhele Director N. Molefe J.F. Thackeray BSc (Hons), MSc (UCT), MPhil, PhD (Yale) A. Mollepolle S. Motsumi Reader L. Sekowe L.R. Berger BA (Hons) (GA Southern), PhD (Witwaters- M. Seshoene rand) Honorary Staff Research Officer Honorary Research Associates K. Carlson BSc (Michigan, Ann Arbor), MSc, PhD Dr Shaw Badenhorst, Transvaal Museum (Indiana, Bloomington) Prof. Steve Churchill, Duke University, USA J. Kibii BSc, (Nairobi), MSc, PhD (Witwatersrand) Prof. Francesco d'Errico, BordeauxUniversity, France B. Kuhn BSc (Washington State), MSc (UCL), PhD (UP) Prof. Daryl de Ruiter, Texas A&M University, USA Research Associates Prof. Katerina Harvati, University of Tubingen, Germany C. Henshilwood BA (Hons) (UCT), PhD (Cambridge) Prof. Jacopo Moggi-Cecchi, Laboratori di Antropologia, M. Lombard BA (Hons), MA (UNISA), PhD (Witwaters- Dipartimento di Biologia Animale e Genetica, Università rand) di Firenze, Italy Administrative and Support Staff Prof. Peter Schmidt, Zurich, Switzerland Senior Administrative Secretary Prof. Himla Soodyall, Human Genomic Diversity and Disease Research Unit (HGDDRU), South African Evlyn Ho Medical Research Council in conjunction with the Preparators National Health Laboratory Service and University of T. Dingiswayo the Witwatersrand PALAEONTOLOGIA AFRICANA ANNALS OF THE BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH UNIVERSITY OF THE WITWATERSRAND ISSN 0078-8554 Volume 44, December 2009 CONTENTS Articles Kemp, T.S. — Phylogenetic interrelationships and pattern of evolution of the 1 therapsids: testing for polytomy Nicolas, M. & Rubidge, B.S. — Assessing content and bias in South African 13 Permo-Triassic Karoo tetrapod fossil collections Weide, D.M., Sidor, C.A., Angielczyk, K.D. & Smith, R.M.H. — A new record 21 of Procynosuchus delaharpeae (Therapsida: Cynodontia) from the Upper Perm- ian Usili Formation, Tanzania Fourie, H. & Rubidge, B.S. — The postcranial skeleton of the basal thero- 27 cephalian Glanosuchus macrops (Scylacosauridae) and comparison of morpho- logical and phylogenetic trends amongst the Theriodontia Govender, R. & Yates,A. — Dicynodont postcrania from the Triassic of Namibia 41 and their implication for the systematics of Kannemeyeriiforme dicynodonts Geraads, D., Melillo, S. & Haile-Selassie, Y. — Middle Pliocene Bovidae from 59 Hominid-bearing sites in the Woranso-Mille area, Afar region, Ethiopia Zipfel, B. & Berger, L.R. — Partial hominin tibia (StW 396) from Sterkfontein, 71 South Africa Technical Note Zipfel, B. & Berger, L.R. — New Cenozoic fossil- bearing site abbreviations for 77 collections of the University of the Witwatersrand Abstracts 15th Biennial Meeting of the Palaeontological Society of Southern Africa •Reviewed Extended Abstracts Abdala, F., Martinelli, A.G., Bento Soares, M., de la Fuente, M. & Ribeiro, A.M. — South American 83 Middle Triassic continental faunas with amniotes: biostratigraphy and correlation Atayman, S., Rubidge, B.S. & Abdala, F. — Taxonomic re-evaluation of tapinocephalid dino- 88 cephalians Backwell, L. & d’Errico, F. — Additional evidence of early hominid bone tools from South Africa. 91 First attempt at exploring inter-site variability Bordy, E.M., Sztanó, O., Rubidge, B.S. & Bumby, A. — Tetrapod burrows in the southwestern main 95 Karoo Basin (Lower Katberg Formation, Beaufort Group), South Africa Browning, C. — Nodular preservation of trilobite fossils from the Bokkeveld Group, Eastern Cape 100 Province, South Africa Manegold, A. — The early fossil record of perching birds (Passeriformes) 103 Mannion, P.D. — Review and analysis of African sauropodomorph dinosaur diversity 108 Matthews, T., Marean, C. & Nilssen, P. — Micromammals from the Middle Stone Age (92–167 ka) at 112 Cave PP13B, Pinnacle Point, south coast, South Africa Mostovski, M.B. — Brachyceran assemblages (Insecta: Diptera) as indicators of terrestrial palaeo- 121 environments in the Late Mesozoic Ovechkina, M.N., Watkeys, M. & Kretzinger, W. — Nannoplankton in the manganese deposits of 126 the Mozambique Ridge and Mozambique Basin, southwestern Indian Ocean Ovechkina, M.N., Watkeys, M. & Mostovski, M.B. — Calcareous nannofossils from the stratotype 129 section of the Upper Cretaceous Mzamba Formation, Eastern Cape, South Africa Rubidge, B. & Angielczyk, K. — Stratigraphic ranges of Tapinocephalus Assemblage Zone dicyno- 134 donts: implications for middle Permian continental biostratigraphy van Dijk, D.E. — Continental displacement: early lines of evidence that deserve attention 136 •Abstracts 139 •Poster Abstracts 190 © 2009 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 Phylogenetic interrelationships and pattern of evolution of the therapsids: testing for polytomy Tom S. Kemp Museum of Natural History and St John’s College, Oxford, OX1 3PW, U.K. E-mail: [email protected] Received 11 October 2008. Accepted 4 May 2009 There is little agreement on the interrelationships of the major therapsid subtaxa because none of the variously proposed sister-group relationships are supported by clearly defined, unambiguously distributed morphological characters. Rather than pursue a new cladistic analysis here, the hypothesis is explored that the lack of an agreed cladogram is because there was a polytomy at the base of the therapsid radiation that is not amenable to positive testing by conventional morphological cladistics, but that can be tested in four ways. The virtually simultaneous appearance of all the lineages except Cynodontia in the Middle Permian stratigraphic record supports the hypothesis. The palaeogeographic record, which shows a combination of taxa with first occurrences in different parts of Pangaea also supports it, though this is not strong evidence. The palaeoenvironmental record supports the polytomous hypothesis strongly by providing evidence of a coincidence between the start of the therapsid radiation and the appearance of a new suite of ecological opportunities for diversification within higher latitudes. Finally, a functional correlation analysis of the characters associated with feeding, and the reconstruction of lineages of functionally integrated organisms offers strong support by indicating that no two of the four respective lineages, Dinocephalia, Gorgonopsia, Anomodontia and Therocephalia, could have shared a functionally feasible common ancestral stage subsequent to a hypothetical ancestor at a biarmosuchian grade. The exception is Cynodontia and Therocephalia, which are inferred to have shared such a more recent common ancestral stage, and therefore to be sister-groups in the taxon Eutheriodonta. Keywords: Therapsida, Permian tetrapods, Permian palaeoecology, correlated progression. INTRODUCTION familiar but at the time utterly unexpected results (e.g. The amniote clade Therapsida is highly significant in the Kemp 2005; Springer et al. 2005). Thus the Therapsida are history of
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