GC University Lahore PALAEOENVIRONMENTAL STUDY OF PAKISTAN SIWALIKS Name: MUHAMMAD TARIQ Session: 2007-2010 Roll No: 05-GCU-PHD-Z-07 Supervisor: Prof. Dr. Nusrat Jahan Department: Zoology PALAEOENVIRONMENTAL STUDY OF PAKISTAN SIWALIKS Submitted to GC University Lahore in fulfillment of the requirements for the award of Degree of Doctor of Philosophy in Zoology by Name: MUHAMMAD TARIQ Session: 2007-2010 Roll No: 05-GCU-PHD-Z-07 Department of Zoology GC University Lahore DEDICATION This work is dedicated to MY LOVING PARENTS (Gulzar Ahmad and Sugran Abdullah) through whom I acquired indefatigable temperament, morality, and determination for achieving objectives in every walk of life MY YOUNGER BROTHERS (Muhammad Yasin and Muhammad Noor-ul-Amin) who gave hope and strength to meet with the challenges of life MY YOUNGER SISTERS Zainab Gulzar, Shahida Gulzar and Shakeela Gulzar for their care, affection and cooperation MY DEAREST DAUGHTER (Kashaf Tariq) who proved great blessing for achieving goals of my life ALL GREAT SOULS who pursue their dreams with persistent efforts to make this world a better place to live in DECLARATION I, Mr. Muhammad Tariq Roll No. 05-GCU-PHD-Z-07 student of PhD in the subject of Zoology session: 2007-2010, hereby declare that the matter printed in the thesis titled “Palaeoenvironmental Study of Pakistan Siwaliks” is my own work and has not been printed, published and submitted as research work, thesis or publication in any form in any University, Research Institution etc in Pakistan or abroad. ________________ _____________________ Dated Signatures of Deponent RESEARCH COMPLETION CERTIFICATE Certified that the research work contained in this thesis titled “Palaeoenvironmental Study of Pakistan Siwaliks” has been carried out and completed by Mr. Muhammad Tariq Roll No. 05-GCU-PHD-Z-07 under my supervision Supervisor: Prof. Dr. Nusrat Jahan Department of Zoology GC University, Lahore Submitted through: Controller of Examinations GC University, Lahore Prof. Dr. Nusrat Jahan Chairperson Department of Zoology GC University, Lahore ACKNOWLEDGEMENTS In the name of Almighty Allah, the most compassionate, ever merciful, who bestowed upon me the wisdom and strength to accomplish this humble piece of research work. I am grateful to my research supervisor Prof. Dr. Nusrat Jahan for her immense support, innovative guidance and patience throughout this research project. I offer my profound thanks to Prof. Dr. Nikos Solounias from New York Institute of Technology, New York, USA for developing collaboration with me regarding microwear and mesowear studies of extinct and extant ungulates. Bundle of thanks to Gina Semprebon and Florent Rivals for sending research literature about microwear studies of ungulates and Mikael Fortelius for appreciating goat experiment on mesowear study and furnishing research literature about it. I am highly thankful to Prof. Christine Janis Marie from Brown University, USA for valuable guidance concerning hypsodonty studies on ungulates and sending masses of research literature. Many thanks to Sherry V. Nelson for her support and sending research literature on Sivapithecus and ungulates from Siwaliks of Pakistan. Best regards to Prof. Dr. John Barry from University of Harvard, USA, for help in estimating the ages of fossil localities of Potwar Siwaliks and sending valuable research papers. Many thanks to the members of departmental academic committee i.e. Dr. Abdul Qayyum Khan Sulehria, Dr. Muhammad Fiaz Qamar and Dr. Iram Liaqat for giving positive suggestions regarding improvement of this dissertation. I am pleased to specially acknowledge Higher Education Commission (HEC) of Pakistan for sponsorship and facilitation of this research project. My special thanks to all the staff members and Ph. D Scholars from Zoology Department, GCU, Lahore, for moral support. I am highly indebted to Raja Jan Muhammad (Late), Raja Tahir Mehmood, Raja Khalid Mehmood and Raja Tariq from Dhok Mori Mothan (Padhri), Anar Khan and Ahmad Shah from Hasnot, Mr. Zakir Hussain from Domeli, Mr. Shahzad from village Ratial, Jhelum District, Muhammad Ajaib, Shukat Ali and Wajid Ali from Dhok Bin MirKhatoon, Chakwal District for their assistance in field work and hospitality. I am thankful to my colleague Muhammad Akram Jan for his help in the field as well as in research work. Kind regards to my younger brothers and sisters for their immense support and cooperation in every aspect of life. MUHAMMAD TARIQ i CONTENTS TOPIC PAGE # ABSTRACT ………………………………………………………………… xi LIST OF ABBREVIATIONS ……………………………………………… x CHAPTER 1 INTRODUCTION…………………………………………………………… 1 1.1.CONSTRUCTION OF CHRONOSTRATIGRAPHIC FRAMEWORK… 5 1.1.1. LITHOSTRATIGRAPHY……………………………………………… 5 1.1.2. BIOSTRATIGRAPHY………………………………………………… 6 1.2. OVERVIEW OF PALEOENVIRONMENT…………………………….. 15 1.3.RESEARCH OBJECTIVES……………………………………………… 20 1.4.SIGNIFICANCE OF WORK……………………………………………… 20 CHAPTER 2 REVIEW OF LITERATURE………………………………………………… 25 CHAPTER 3 MATERIALS AND METHODS………………………………………………… 32 3.1. MESOWEAR TYPE I&II METHOD……………………………………… 32 3.2. MESOWEAR TYPE III METHOD………………………………………… 33 3.2.1. DESCRIPTION OF METHOD…………………………………… 33 3.2.2. DEVELOPMENT OF SCORING SCALES...…………………… 34 3.2.2.1. BAND-2 SCORING SCALE…………………………………… 34 3.2.2.2. J-POINT SCORING SCALE…………………………………… 35 3.3. MICROWEAR METHOD………………………………………………… 36 3.3.1. ESTABLISHING MORPHOSPACES……………………………… 37 3.3.2. QUANTITATIVE CHARATERIZATION OF AVERAGE MICROWEAR COUNTS………………………………………… 38 3.4. HYPSODONTY METHOD………………………………………… 38 3.4.1. DIETARY AND HABITAT CHARACTERIZATION OF SIWALIK MAMMALS………………………………………… 39 3.4.2. MEAN HYPODONTY………………………………………… 41 ii CHAPTER 4 RESULTS..…………………………………………………………… 54 4.1. MESOWEAR TYPE-1 AND II ANALYSES…………………… 54 4.1.1. Giraffokeryx punjabiensis…………………………………… 54 4.1.2. Tragoportax sp…………………………………………… 54 4.1.3. Selenoportax sp………………………………………………… 55 4.1.4. Gazella lydekkeri……………………………………………… 55 4.2. MESOWEAR TYPE III ANALYSIS……………………………… 55 4.3. MICROWEAR ANALYSIS………………………………………… 57 4.3.1. DIETARY ADAPTATIONS OF UNGULATES BASED ON MICROWEAR................................................................ 57 4.3.1.1. Suidae…………………………………………………… 57 4.3.1.2. Tragulidae…………………………………………………… 58 4.3.1.3. Boselaphini…………………………………………………… 58 4.3.1.4. Giraffidae……………………………………………………… 59 4.3.1.5. Anthracotheriidae……………………………………………… 59 4.3.1.6. Hiparion sp…………………………………………………… 59 4.3.1.7. Rhinocerotidae……………………………………………….. 59 4.4. HYPSODONTY ANALYSIS……………………………………… 59 4.4.1. Boselaphini…………………………………………………… 59 4.4.2. Bovini………………………………………………………… 60 4.4.3. Reduncini and Antilopini...…………………………………… 60 4.4.4. Giraffids……………………………………………………….. 61 4.4.5. Tragulidae……………………………………………………… 61 4.4.6. Cervidae………………………………………………………… 62 4.4.7 Suidae…………………………………………………………… 62 4.4.8 Equidae………………………………………………………. 62 4.4.9. Rhinocerotidae……………………………………………….. 62 4.5. MEAN HYPSODONTY ANALYSIS………………………………… 63 iii CHAPTER 5 DISCUSSION……………………………………………………………… 106 5.1. PALEOCLIMATIC RECONSTRUCTION…………………………… 106 5.1.1. IMPACT OF PALEOCLIMATE ON FAUNAL EVOLUTION…… 114 5.2. EXPLORATION OF PALEOVEGETATION..…………………………… 119 5.2.1. DRIVERS FOR CHANGE FROM FOREST TO C3-C4 TRANSITION………………………………………. 128 5.2.2. RATIONAL FOR DOMINANCE OF C4 GRASSLANDS………… 129 5.3. SPECIES DIVERSITY ………………….………………………………… 132 5.3.1. SPECIES RICHNESS IN SIWALIK UNGULATES……………… 132 5.3.2. RODENT (MURID-CRICETID) SPECIES RICHNESS (DIVERSITY) THROUGH THE MIO-PLIOCENE IN THE SIWALIKS…………………………………………….. 137 5.3.3. FAUNAL TURNOVER, RELATIVE ABUNDANCE AND CHANGE OF PALEOCOMMUNITY STRUCTURE…… 138 REFERENCES…………………………………………………………………… 160 APPENDICES…………………………………………………………………… 189 iv LIST OF TABLES RESULTS Table 4.1; Absolute and relative mesowear scorings of upper M2s of Giraffokeryx punjabiensis………………………………… 64 Table 4.2; Absolute and relative mesowear scorings of upper M2s of Tragoportax sp…………………………………………… 64 Table 4.3; Absolute and relative mesowear scorings of upper M2s of Selenoportax,sp. ……………………………………… 64 Table 4.4; Absolute and relative mesowear scorings of upper M2s of Gazella lydekkeri …………………………………… 64 Table 4.5; Data for Mesowear II and III for Extant Ruminants……… 65 Table 4.6; Data for Mesowear II and III for Extinct species of ungulates from Siwaliks of Pakistan and of Giraffids from late Miocene of China……………………………………………… 66 Table 4.7; Fossil species from late Miocene of Pakistan Siwaliks and their dental microwear results………………………… 67 Table 4.8; Ecomorphic data of boselaphines of Siwaliks of Pakistan…………………………………………… 68 Table 4.9; Ecomorphic data of antilopines of Siwaliks…………………….. 69 Table 4.10; Ecomorphic data of boviines of Siwaliks………………………. 69 Table 4.11; Ecomorphic data of giraffids of Siwaliks………………………. 70 Table 4.12; Ecomorphic data of Siwalik tragulids…………………………… 70 Table 4.13; Ecomorphic data of cervids of Siwaliks……………………….... 71 Table 4.14; Ecomorphic data of suids of Siwaliks…………………………… 71 Table 4.15; Ecomorphic data of Siwalik horses…………… ……………… 72 Table 4.16; Ecomorphic data of Siwalik rhinoceroses……………………… 72 Table 4.17; Mean hypsodonty values of major localities included in the study… 73 Table 4.18; Dietary characterization in extict giraffids based on p-values for the sample……………………………………………. 75 Table 5.1; Infrences of mean annual precipitation from mean ordinate hypsodonty values ……………………………………… 149 v Table 5.2; Species richness in ungulates recorded from Siwaliks of Pakistan… 150 Table 5.3; Relative abundance among different ungulate groups from Pakistan Siwaliks………………………………………………… 152 Table 5.4; Summary of evolution of ungulate