Miocene rodent evolution and migration Muroidea from Pakistan, Turkey and Northern Africa W. Wessels GEOLOGICA ULTRAIECTINA Mededelingen van de Faculteit Geowetenschappen departement Aardwetenschappen Universiteit Utrecht No. 307 ISBN 978-90-5744-170-7 Graphic design and figures: GeoMedia, Faculty of Geosciences, Utrecht University (7497) Miocene rodent evolution and migration Muroidea from Pakistan, Turkey and Northern Africa Evolutie en migratie van Miocene knaagdieren Muroidea afkomstig uit Pakistan, Turkije en Noord Afrika (met een samenvatting in het Nederlands) PROEFSCHRIFT ter verkrijging van de graad van doctor aan de Universiteit Utrecht op gezag van de rector magnificus, prof.dr. J.C. Stoof, ingevolge het besluit van het college voor promoties in het openbaar te verdedigen op maandag 8 juni 2009 des middags te 4.15 uur door Wilma Wessels geboren op 9 december 1955 te Vriezenveen Promotor: Prof.dr. J.W.F. Reumer Contents Part 1 Introduction 1 Introduction 13 2 Correlation of some Miocene faunas from Northern Africa, Turkey and Pakistan 17 by means of Myocricetodontidae Published in Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen B 90(1): 65-82 (1987), Wessels W., Ünay. E. & Tobien H. 2.1 Abstract 17 2.2 Introduction 17 2.3 The Pakistani Myocricetodontinae 19 2.3.2 Taxonomy 19 2.3.3 Discussion of the Pakistani Myocricetodontinae 24 2.4 The Turkish Myocricetodontinae 24 2.4.2 Taxonomy 25 2.4.3 Discussion of the Turkish Myocricetodontinae 31 2.5 Conclusions 31 2.6 Acknowledgements 31 Part 2 Rodents from Europe, Turkey and Northern Africa 3 Gerbillidae from the Miocene and Pliocene of Europe 35 Published in Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie 38: 187-207 (1998), Wessels W. 3.1 Abstract 35 3.2 Introduction 35 3.3 The Family Gerbillidae 35 3.4 Taxonomy 37 3.4.1 Introduction 37 3.4.2 Subfamily Myocricetodontinae 39 3.4.3 Subfamily Taterillinae 41 3.4.4 Subfamily Gerbillinae 43 3.5 Migrations of the European Gerbillidae 46 3.6 Acknowledgements 46 Notes 46 Appendix 3.1 Occurrences of Gerbillidae in Europe 47 Appendix 3.2 Fossil occurrences of Gerbillidae in Europe, Africa and Asia 48 5 4 Miocene small mammals from Jebel Zelten, Libya 51 Published in Miocene small mammals from Jebel Zelten, Libya. In: López-Martínez N., Peláez-Campomanes P. & Hernández Fernández M. (Eds.), Surrounding Fossil Mammals: Dating, Evolution and Paleoenvironment. Coloquios de Paleontología, Volumen Extraordinario 1, en homenaje al Dr. Remmert Daams: 699-715 (2003), Wessels W., Fejfar O., Peláez-Campomanes P., Meulen A.J. van der & Bruijn de H. 4.1 Abstract 51 4.2 Introduction 52 4.3 Taxonomy 53 4.4 Discussion and conclusions 70 4.5 Acknowledgements 72 5 Myocricetodontinae and Megacricetodontini from the lower Miocene of NW Anatolia 73 Published in Myocricetodontinae and Megacricetodontini (Rodentia) from the lower Miocene of NW Anatolia. In: Horáček I. & Míkovský J. (Eds.), Papers in paleomammalogy honoring Prof. Dr. Oldřich Fejfar. Lynx N.S. 32: 371-388(2001), Wessels W., Theocharopoulos K.D., Bruijn H. de & Ünay E. 5.1 Abstract 73 5.2 Introduction 73 5.3 Methods 75 5.4. The subfamily Myocricetodontinae Lavocat, 1961 75 5.4.1 Introduction 75 5.4.2 Systematic paleontology 75 5.4.3 Description and discussion of the skull and mandible fragments 82 5.5 The subfamily Cricetinae s.l. Fahlbusch, 1964 83 5.5.1 Systematic paleontology 83 5.5.2 Discussion of Megacricetodon 85 5.6 Discussion of Vallaris and Megacricetodon 86 5.7 The Keseköy assemblage 86 5.8 Acknowledgements 87 6 Progonomys from the Kütahya area (Turkey) 89 6.1 Introduction 89 6.2 Material and methods 90 6.2.1 Material 90 6.2.2 Measurements 92 6.2.3 Amount of material 92 6.2.4 Description and nomenclature 92 6.2.5 Species concept 93 6.2.6 Comparison material 97 6.3 Taxonomy 97 6.3.1 Material from Altıntaş and Kütahya 97 6.3.2 Progonomys cathalai Schaub, 1938 97 6.3.2.1 Description 97 6.3.2.2 Comparison and discussion 106 6.4 Overview of Vallesian and early Turolian murines 107 6 6.5 Discussion on the genus Progonomys Schaub, 1938 110 6.5.1 Definition 110 6.5.2 Discussion 111 6.5.3 Differences and affinities of Progonomys 112 6.5.4 Progonomys species 113 6.5.5 Distribution of Progonomys 113 6.6 Discussion and conclusions 117 Appendix 6.1 119 Part 3 Rodents from Pakistan 127 7 Myocricetodontinae from the Miocene of Pakistan 129 Published in Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen C 99(3-4): 253-312 (1996), Wessels W. 7.1 Abstract 129 7.2. Introduction 129 7.3 Taxonomy 132 7.3.1 Superfamily Muroidea Miller & Gidley, 1918 132 7.3.2 Family Myocricetodontidae 132 7.4 Conclusions 176 7.5 Acknowledgements 180 8 Rhizomyidae from the Lower Manchar Formation (Miocene, Pakistan) 183 Published in Annals of Carnegie Museum 70(2): 143-168 (2001), Wessels W. & Bruijn H. de 8.1 Abstract 183 8.2 Introduction 183 8.3 Systematic paleontology 184 8.3.1 Introduction to the Family Rhizomyidae Miller & Gidley, 1918 184 8.3.2 Taxonomy 185 8.4 The Miocene record of the Rhizomyidae in Pakistan 197 8.5 The potential of the Rhizomyidae for biostratigraphy 198 8.6 The age of the Gaj and Sehwan assemblages 198 8.7 Acknowledgments 199 Appendix 8.1 200 Appendix 8.2 201 Appendix 8.3 202 Appendix 8.4 203 Appendix 8.5 204 Appendix 8.6 204 Appendix 8.7 205 Appendix 8.8 206 Appendix 8.9 206 Appendix 8.10 207 Appendix 8.11 207 7 9 Murinae from the Lower Manchar Formation, Pakistan 209 9.1 Introduction 209 9.2 Taxonomy 210 9.2.1 Potwarmus 210 9.2.2 Antemus 215 9.2.3 Comparison and discussion of Antemus mancharensis 220 9.2.4 Wear patterns and chewing directions 222 9.4 Conclusions 223 9.5 Acknowledgements 225 Part 4 Synthesis and outlook 227 Miocene rodents, faunal exchange and migrations routes between Eurasia and Africa 229 10.1 Introduction 229 10.2 The fossil rodents from Pakistan 231 10.2.1 The rodents from the Lower Manchar Formation 231 10.2.2 Constraints on dating the Sehwan and Gaj assemblages 232 10.2.3 Rodent occurrences in Pakistan 233 10.2.4 Climate change and rodent development in Pakistan 234 10.3 Fossil rodents from Northern Africa, the Arabian Plate and Anatolia 235 10.3.1 Northern Africa 235 10.3.2 Arabian Plate 235 10.3.3 Anatolia 235 10.4 Overview of taxonomy and phylogeny 236 10.4.1 Rhizomyinae 236 10.4.2 Myocricetodontinae 236 10.4.3 Murinae 238 10.4.4 Ctenodactylidae 238 10.4.5 Other rodent groups 239 10.5 Selected data on large mammals 241 10.5.1 The biostratigraphical position of the “Bugti Fauna” 241 10.5.2 Data on the distribution of African large mammals 242 10.5.3 Data on the distribution of European large mammals 243 10.6 Faunal exchange and migration routes 243 10.6.1 Introduction 243 10.6.2 Plate tectonics and palaeogeographic settings of the Arabian Plate and 244 the Tethys 10.6.3 Plate tectonics and palaeogeographic settings of the Indian plate 245 10.6.4 Possible mammal migration routes 245 10.6.5 Faunal exchange 247 Appendix 10.1 Distribution chart with numbers of all molar 251 Appendix 10.2 Distribution chart with numbers of all M1 and M2 252 8 References 253 Summary 279 Samenvatting 283 Dankwoord 287 Curriculum Vitae 288 List of Publications 289 9 10 Part 1 Introduction 11 12 CHaptER 1 Introduction The distribution of terrestrial and marine fauna and flora in space and time gives us a glimpse of the complexity of life on earth. However, the amount and variability of organisms involved is tremendous: literally millions of organisms at millions of localities. To order this vast amount of information we need tools. Biostratigraphy is such an essential tool; with biostratigraphical methods we correlate sedimentary rocks containing fossils, over short or long distances, and assign relative ages. In combination with other dating methods, such as magnetostratigraphy and isotopic dating, absolute ages can be assigned. Together these methods build and order the puzzle of life through time. Taxonomy is essential in biostratigraphy; species recognition and classification is the first step in evolution interpretation and the reconstruction of migration events. Mammals can be used in correlating and dating in the terrestrial realm. Rodents are especially suited, since they have a short life span and high productivity, and further adapt rapidly to changing environments. Their dental elements fossilize very well and, therefore, their evolutionary development can be followed relatively easily in the fossil record. In this thesis, molars of fossil rodents are used as a biostratigraphical tool in correlating and dating Pakistani, Turkish and Libyan fossil assemblages. In many assemblages the Muridae (superfamily Muroidea), including the subfamilies of the Myocricetodontinae, Murinae and Rhizomyinae, is the most dominant rodent group present. The taxonomy, evolutionary histories and distributions of the many Muridae species in time and space are extensively described and discussed. The lithostratigraphy of the Pakistani and Libyan sequences is described in Chapters 4 and 7, respectively. Lithostratigraphic positions of some of the Turkish assemblages can be found in Chapters 5 and 6. The nomenclature used for the dental structures of the Murinae is discussed in Chapter 6, for the Myocricetodontinae and Rhizomyinae such a nomenclature is provided in Chapter 7. Not only fossil rodents document changes in time. From the first published paper to the more recent ones, this thesis reflects changes in the taxonomical hierarchy used, and in the way molars are described.