THE Evolution and Palaeoecology of Elephants and Their Relatives

Edited by JEHESKEL SHOSHANI , , USA and Elephant Research Foundation, Bloomfield Hills, Michigan, USA and PASCAL TASSY URA 12, CNRS, Universite Paris VI, Laboratoire de Paleontologie des Vertebres et de Paleontologie Humaine, Paris, France

Oxford New York Tokyo OXFORD UNIVERSITY PRESS 1996 CONTENTS

Foreword ix

VINCENT J. MAGLIO

List of contributors xix Henry Fairfield Osborn and the Proboscidea xxiii EDWIN H. COLBERT List of abbreviations, symbols, and terms xxix

PART I INTRODUCTION

1. Historical overview of classification and phylogeny of the Proboscidea 3 PASCAL TASSY AND JEHESKEL SHOSHANI 2. Skeletal and other basic anatomical features of elephants 9 JEHESKEL SHOSHANI 3. Dental homologies and nomenclature in the Proboscidea 21 PASCAL TASSY 4. Ontogenetic evidence for incisor homologies in proboscideans 26 W. PATRICK LUCKETT

PART II PROBOSCIDEAN RELATIVES

5. On the position of Proboscidea in the phylogenetic system of Eutheria: a systematic review 3 5 MARTIN S. FISCHER 6. Who is who among the Proboscidea? 39 PASCAL TASSY 7. Proboscidean relationships based on immunological data 49 JEROLD M. LOWENSTEIN AND JEHESKEL SHOSHANI

PART III THE FIRST RADIATION: EARLY PROBOSCIDEANS

8. The earliest proboscideans: general plan, , and palaeoecology 5 7 JEHESKEL SHOSHANI, ROBERT M. WEST, NICHOLAS COURT, ROBERT J. G. SAVAGE, and JOHN M. HARRIS 9. Evolution of zygodons with emphasis on dentition 76 HEINZ TOBIEN CONTENTS PART IV THE SECOND RADIATION: AND STEGODONTIDS

10. The earliest gomphotheres 89 PASCAL TASSY 11. Growth and sexual dimorphism among Miocene elephantoids: the example of Gomphotherium angustidens 92 PASCAL TASSY 12 A. Phylogeny of African and Eurasian Elephantoidea of the late Neogene 101 JON E. KALB, DAVID J. FROEHLICH, and GORDEN L. BELL 12B. Palaeobiogeography of late Neogene African and Eurasian Elephantoidea 117 JON E. KALB, DAVID J. FROEHLICH, and GORDEN L. BELL 13. On the shovel-tusked elephantoids from China 124 JIAN GUAN 14. Gomphotheres and mammutids from the Iberian Peninsula 136 ANA V. MAZO 15. The biogeography of the gomphotheriid proboscideans of North America 143 W. DAVID LAMBERT 16. Para- or monophyly of the gomphotheres and their position within Proboscidea 149 JEHESKEL SHOSHANI 17. Stegodontidae: evolutionary relationships 178 HARUO SAEGUSA

PART V THE THIRD RADIATION:

18. Origin and radiation of the Elephantidae 193 NANCY E. TODD and V. LOUISE ROTH 19. Evolution and taxonomy of Eurasian mammoths 203 ADRIAN M. LISTER 20. Plio-Pleistocene proboscideans from the southern bight of the North Sea and the Eastern Scheldt, The Netherlands 214 HANS VAN ESSEN and DICK MOL 21. Proboscidean remains in southern Levant 225 EITAN TCHERNOV and JEHESKEL SHOSHANI 22. The Pleistocene dwarf elephants of Mediterranean islands 234 LUCIA CALOI, TASSOS KOTSAKIS, MARIA R. PALOMBO, and CARMELO PETRONIO 23. The proboscideans of the South-East Asian islands 240 GERT D. VAN DEN BERGH, PAUL Y. SONDAAR, JOHN DE VOS, and FACHROEL AZIZ 24. Pleistocene dwarf elephants of the California Islands 249 V. LOUISE ROTH CONTENTS 25. Sexual dimorphism in the mammoth pelvis: an aid to gender determination 254 ADRIAN M. LISTER 26. Sexual dimorphism in the mammoth skull, teeth, and long bones 260 ALEXANDER 0. AVERIANOV

PART VI PALAEOECOLOGY, TAPHONOMY, EXTINCTION, AND CONSERVATION

27. North American Mammutidae 271 JEFFREY J. SAUNDERS 28. Distribution and palaeoecology of central and western North American Mammuthus 280 LARRY D. AGENBROAD and JIM I. MEAD 29. Mammoths, gomphotheres, and the Great American Faunal Interchange 289 JOSEPH P. DUDLEY 30. Extinction of proboscideans in North America 296 DANIEL C. FISHER 31. South American proboscideans: general introduction and reflections on Pleistocene extinctions 316 RODOLFO M. CASAMIQUELA, JEHESKEL SHOSHANI, and THOMAS D. DILLEHAY 32. Loxodonta africana: range and distribution, past and present 321 IAIN DOUGLAS-HAMILTON and FRANCES MICHELMORE 33. maximus: status and distribution 327 RAMAN SUKUMAR and CHARLES SANTIAPILLAI

PART VII SUMMARY AND CONCLUSIONS

34. Summary, conclusions, and a glimpse into the future 335 JEHESKEL SHOSHANI and PASCAL TASSY

APPENDICES

A Synonyms of proboscidean taxa, genera and subgenera 349 B A suggested classification of the Proboscidea 352 C A summary of proboscidean taxa 354 D Geological time-scale and correlation chart among Land Mammal Ages 387

REFERENCES 391 AUTHOR INDEX 433 STRATIGRAPHIC, GEOGRAPHIC, AND LOCALITY INDEX 441 TAXA INDEX 450 SUBJECT INDEX 462