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Cranial Morphology of a Primitive Dinocephalian from The CRANIAL MORPHOLOGY OF A PRIMITIVE DINOCEPHALIAN FROM THE MADUMABISA MUDSTONE FORMATION, ZIMBABWE. Darlington Munyikwa A dissertation submitted to the Faculty of Science, University of Witwatersrand, Johannesburg, in fulfilment of the requirements for the degree of Master of Science. Johannesburg 2001 ii DEC LARATION I declare that this dissertation is my own, unaided work except where it is indicated in the text. This dissertation is submitted for the degree of Master of Science at the University of Witwatersrand, Johannesburg. It has not been submitted before for any degree or any other qualification at any other university. (Signature of candidate) ;"4 day of_-'-M----'------'-t7-_j~ ___ 2001. iii ABSTRACT Dinocephalians form an important part of the Upper Permian therapsid faunas of South Africa and Russia, and have also recently been reported from China and Brazil, and even more recently a diverse dinocephalian fauna has been described from Zimbabwe. This thesis reports a new primitive tapinocephalid dinocephalian, NHMB 1556, from the Upper Madumabisa Mudstone Formation in Zimbabwe. NHMB 1556 is considered to be closely related to Avenantia and it possesses primitive tapinocephalid characters, but is more derived than Tapinocaninus, the most primitive tapinocephalid dinocephalian known. NHMB 1556 is distinguished from other tapinocephalines by having a groove on the squamosal below its dorsal contact with the parietal, a low squamosal-parietal suture on the posterior border of the temporal fenestra, a vomer which extends posteroventrally and forms the anterior margin of the interpterygoidal vacuity and basisphenoid, which is semi-circular anteroventrally. iv TO MY CHILDREN v ACKNOWLEDGEMENTS My most sincere thanks go to my supervisors Prof. B.S. Rubidge and Dr. M.A. Raath for assisting me in selecting the appropriate project for study under the prevailing circumstances and for directing me throughout the entire program of study. I am very grateful to my supervisors for their constructive comments, encouragement, and moral and material support. Their scholarship, academic reputation, and love of the subject were constant sources of inspiration to me. I am very grateful to the South Africa Museum (SAM) for allowing me unlimited access to their collection. Great thanks go to the SAM staff in the Division of Earth Sciences, particularly Dr R.M.H. Smith and S. Kaal, for assisting me in the collection and for their hospitality during my two visits. The assistance and encouragement received from the staff at the Bernard Price Institute for Palaeontological Research (BPI) are greatly appreciated. Dr. S.P. Modesto and Dr. R. ]. Damiani were very helpful with the phylogenetic analysis and guidance in preparation of descriptive figures. I am indebted to the Board Members and the Directorate of National Museums and Monuments of Zimbabwe (NMMZ) for granting me study leave enabling me to visit the vi BPI and SAM. I would like also to thank NMMl staff Messrs L. Ngwenya, L. londo and M. londo for their assistance during the preparation of the specimen. I am also thankful to Mr T. Gumbi, a former employee ofthe Wankie Colliery Company, for bringing to the attention of Natural History Museum, Bulawayo (NHMB), the discovery of the Hwange beds that bear the therapsid fossils. The Wankie Colliery Company made it possible for us to carry out the research by granting us unconditional access to the fossil sites as well as their kind co-operation and hospitality during our visits. All efforts by several geologists and palaeontologists, including Dr. 1. Lepper, B. Barber, Prof. G. Bond, Dr M.A. Raath, Prof. B.S. Rubidge, to unravel the geology and palaeontology of the Madumabisa Mudstones at Hwange, are greatly appreciated. NMMl, John Wakeford Scholarship Trust and myself jointly sponsored the study program. I am therefore indebted to the two funding institutions for without their financial support I could not have been able to complete the study program. I would like also to thank Prof. Bruce S. Rubidge for securing funds for my accommodation during my last visit at BPI and to attend the Palaeontological Society of Southern Africa Conference, Pretoria. Last, but not least, I would like to thank my family for graciously releasing me from most of my daily responsibilities in my quest for a higher education. Their tolerance and vii support are greatly appreciated. I am dedicating my thesis to my two children, Tariro Clarity and Taridzai Brian. viii TABLE OF CONTENTS DECLARATION II ABSTRACT 111 ACKNOWLEDGEMENTS v LIST OF TABLES x LIST OF FIGURES x LIST OF APPENDIXES XI ABBREVIA TIONS USED IN THE FIGURES XII 1 INTRODUCTION 1.1 Therapsid origins 1.2 Historical review of the dinocephalian taxonomy 4 2 GEOLOGICAL BACKGROUND TO THE STUDY AREA 18 2.1 Dwyka 20 2.2 Ecca 20 2.3 Madumabisa Mudstone 22 ix 3 MATERIALS AND METHODS 26 3.1 Materials 26 3.2 Methods 28 3.2.1 Specimen preparation 28 3.2.2 Phylogenetic analysis 30 4 DESCRIPTION OF SKULL 33 4.1 Skull roof 34 4.2 Occiput 42 4.3 Palate 46 4.4 Braincase 50 5 DISCUSSION 52 6 PHYLOGENETIC ANALYSIS 58 7 CONCLUSION 62 8 REFERENCES 64 x TABLES Tablel. Characteristic features of dinocephalian families 8 Table 2. Characters of the subfamilies of the Tapinocephalidae 9 Table 3 Classification of the Dinocephal ia 10 Table 4. Characters of the tribes of the Tapinocephalinae 13 ILLUSTRATIONS Figure l. Cladogram of the families of the Dinocephalia I I Figure 2. Cladogram of the main groups of the Dinocephalia 12 Figure 3. Cladograms of the Dinocephalia with the inclusion 15 of the Stracocephalidae and illustrating Anteosauridae as the most basal group Figure 4. Locality map of the study area 19 Figure 5. Stratigraphic section showing the horizon of 21 dinocephalian fossils xi Figure 6. Some of the better preserved dinocephalian 29 fossils from Hwange Figure 7. Dorsal view ofNHMB1556 35 Figure 8. Lateral view ofNHMB 1556 37 Figure 9. Occipital view ofNHMB 1556 43 Figure 10. Ventral view ofNHMB1556 47 Figure 11. Cladogram illustrating interrelationships 60 of dinocephalian genera APPENDIXES Appendix I. List of characters used in the cladistic analysis Appendix 2. Character matrix xii ABBREVIATIONS USED IN THE TEXT FIGURES boc basioccipital bs basisphenoid earn external auditory meatus ect ectopterygoid f frontal fjug foramen jugularis fmag foramen magnum fo fenestra ovalis iptv interpterygoidal vacuity J jugal lpt lateral process of pterygoid mx maxilla n nasal op opisthotic p parietal pal palatine pf pineal foramen xiii pm premaxilla po postorbital pof postfrontal pp postparietal prf prefrontal prt prootic ptf posttemporal fenestra q quadrate q] quadratojugal sq squamosal t tabular vo vomer CHAPTER 1: INTRODUCTION 1.1 THERAPSID ORIGINS The Dinocephalia, together with the clades Biarmosuchia, Eotitanosuchia, Anomodontia (including Dicynodontia), Gorgonopsia, Therocephalia and Cynodontia constitute the Therapsida (Hopson 1991; Hopson and Barghusen 1986; Hotton 1991; Kemp 1988). Therapsids are considered to have arisen from sphenacodont "pelycosaurs" during the early Late Permian and to be the lineage leading to mammals (Hopson 1967, 1991; Hopson and Barghusen 1986; Kemp 1982; Kermack and Kermack 1984; Rowe 1988; Sidor and Hopson 1998). A great deal of research on therapsid origin, diversification and biogeography has been undertaken and their remains have been found on every continent (Anderson and Cruickshank 1978; Colbert 1986; Hopson 1991; Hopson and Barghusen 1986; Kemp 1982, 1988; King 1988; Olson 1986; Parrish, Parrish and Ziegler 1986; Romer 1970; Sigogneau­ Russell 1989). Biarmosuchids and eotitanosuchids are considered to be the most basal therapsids (Hopson 1991; Hopson and Barghusen 1986; Kemp 1988). Most of the advanced therapsids have a wide distribution and have a well-preserved fossil record. In contrast, the basal therapsids have a limited distribution and a poor fossil record and hence their evolutionary history is vague as compared with their descendants. The earliest therapsid faunas include biarmosuchids, eotitanosuchids, dinocephalians, non- 2 dicynodont anomodonts, gorgonopsians and therocephalians, and are known only from South Africa (Boonstra 1963a, 1963b, 1969; Brinkman 1981; King 1988; Modesto, Rubidge and WeIman 1999; Modesto and Rubidge 2000; Rubidge, Kitching and van den Heever 1983; Rubidge and Hopson 1990, 1996; Rubidge 1991, 1994), Russia (Ivakhnenko 1994, 1996; Olson 1962; Sigogneau and Chudinov 1972) China (Cheng and Li 1997; Li, Rubidge and Cheng 1996), Zimbabwe (Bond 1973; Boonstra 1946) and Brazil (Langer 1998). Olson (1962, 1986) gave a detailed account of the Late Permian terrestrial vertebrates of the San Angelo Formation of the United States of America in which he erected the therapsid infraorders Eotheriodontia and Eutheriodontia ofthe suborder Theriodontia, Eodinocephalia and Eudinocephalia of the suborder Dinocephalia, and Venjukovioidea of the suborder Anomodontia. Parrish et al. (1986) recognised Olson's early therapsids from North America but because of poor preservation he doubted their taxonomic affinities. Recent examination by Sidor and Hopson (1995) has confirmed their "pelycosaurian" status. Dinocephalians, anomodonts, gorgonopsians and therocephalians have been found in the Lower Beaufort Group of South Africa (Boonstra 1963a, 1963b, 1969, 1971; Brinkman 1981; King 1988; Kitching 1977; Modesto c( af. 1999; Rubidge 1987, 1988, 1991, 1994 1995; Rubidge and Hopson 1990, 1996; Rubidge et af. 1983; Smith and Keyser 1995). In addition to dinocephalians, anomodonts,
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