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
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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
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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
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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 community in Pakistan Siwaliks………………………………………… 154 LIST OF FIGURES Figure 1.1; Map showing geographic placement of Siwalik Hills 22 Figure1.2; Map of Potwar Siwaliks displaying major geological formations 22 Figure 1.3; Locality map of Pabbi Hills (Pinjore Formation), district Gujrat, Pakistan 23 Figure 1.4; Biochronostratigraphic framework of Siwaliks of Pakistan and its correlation with European and Chinese chronology…………… 24 Figure 3.1; Tooth mesowear type-1displaying selection of three types of apices… 42 Figure 3.2; The concept of Mesowear-I based on two variable named cusp relief and cusp shape (sharp, round and blunt).. …………… 43 Figure 3.3 A; Mesowear “ruler” designed for dental mesowear II scorings…… 44 Figure 3.3 B; The mesowear “ruler” applied to a fossil bovid- Urmiatherium……………………………………………….. 44 Figure 3.4; Pattern of mesowear-III of a typical browser…………………… 45 Figure 3.5; Pattern of mesowear-III of a typical grazer……………………… 46 Figure 3.6; A model made in clay showing coding system of mesowear-III…………………………………… 47 Figure 3.7A; Mesowear-III of browsing goats; B10 and B20………………… 48 Figure 3.7B; Mesowear- III of browsing goats; B30 and B40. ……………… 49 Figure 3.8A; Mesowear- III of grazing goats; G10 and G20…………………… 50 Figure 3.8B; Mesowear- III of grazing goats; G30 and G40. ………………… 51 Figure 3.9 A-E; A. Mammalian molar with demarcated second band of paracone.. where all microwear molds and cast preparation is focused. ……… 52 Figure 3.10; Generalized SEM photomicrographs at 500X. …………………… 53 Figure 4.1; Mesowear scorings displaying Cusp relief and cusp shape of M2 of Giraffokeryx punjabiensis…………………………………… 76 vi
Figure 4.2; Dendrogram showing the hierarchical cluster analyses of Giraffokeryx punjabiensis……………………………………… 76 Figure 4.3 (A, B, C); Bivariate plots of hypsodonty index (HI) versus %age of high occlusal relief. ……………………………………… 77 Figure 4.4 (A, B, C); Mesowear analyses of cusp shape in G.punjabiensis……. 78 Figure 4.5 (A, B, C); Mesowear analyses of cusp shape in G. punjabiensis … 79 Figure 4.6 (A, B, C); Mesowear analyses of cusp shape in G. punjabiensis……… 80 Figure 4.7 (A, B, C); Mesowear scorings displaying cusp relief, cusp shape of M2 of Tragoportax sp…………………………………… 81 Figure 4.8; Dendrogram showing the hierarchical cluster analyses of Tragoportax sp...... 81 Figure 4.9 (A, B, C); Bivariate plots of hypsodonty index (HI) versus percentage of high occlusal relief…………………………… 82 Figure 4.10; Mesowear scorings displaying Cusp relief and cusp shape of M2 of Selenoportax sp. ………………………………………… 83 Figure 4.11; Dendrogram showing the hierarchical cluster analyses of Selenoportax sp……………………………………………… 83 Figure 4.12 (A, B, C); Bivariate plots of hypsodonty index (HI) versus percentage of high occlusal relief……………………………… 84 Figure 4.13; Mesowear scorings displaying cusp relief and cusp shape of M2 of Gazella lydekkeri………………………………………… 85 Figure. 4.14; Dendrogram showing the hierarchical cluster analyses of Gazella lydekkeri ……………………………………………… 85 Figure 4.15 (A, B, C); Bivariate plots of hypsodonty index (HI) versus percentage of high occlusal relief………………………………… 86 Figure 4.16 (a, b); shows the separation between the extant wild grazers and browsers using a 2 sample t-test assuming unequal variance… 87 Figure 4.17; Results for the goat experiment…………………………………….… 88 Figure 4.18; Rating of fossil species of Giraffids from Siwaliks and Late Miocene of China. ………………………………………… 88 Figure 4.19; Dendrogram showing the hierarchical cluster analyses vii
of E. khauristanensis…………………………………………… 89 Figure 4.20; Dendrogram showing the hierarchical cluster analyses of Dorcatherium majus…………………………………………… 90 Figure 4.21; Dendrogram showing the hierarchical cluster analysis of Giraffokeryx punjabiensis………………………………………… 90 Figure 4.22; Dendrogram showing the hierarchical cluster analysis of Hipparion sp…………………………………………………… 91 Figure 4.23; Dendrogram showing the hierarchical cluster analysis of Merycopotamus nanus…………………………………………… 91 Figure 4.24; Dendrogram showing the hierarchical cluster analysis of Gazella lydekkeri…………………………………………… 92 Figure 4.25; Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and fossil suids…..… 93 Figure 4.26; Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and fossil tragulids.…… 94 Figure 4.27; Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and boselaphine remains…………………………………………………………… 95 Figure 4.28; Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and fossil Giraffids……………………………………………… 96 Figure 4.29; Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and fossil of anthracothroides…………………………………………………….97 Figure 4.30; Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and fossil equids... 98 Figure 4.31; Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and fossils of rhinos……………………………………………… 99 Figure 4.32; Hypsodonty based ecomorphic information of boselaphine remains.. 100 Figure 4.33; Hypsodonty based ecomorphic information of bovine remains ….. 100 viii
Figure 4.34; Hypsodonty based ecomorphic information of fossils of antilopes…. 101 Figure 4.35; Hypsodonty based ecomorphic information of Sivatherine remains… 101 Figure 4.36; Hypsodonty based ecomorphic information of fossils of giraffinae…. 102 Figure 4.37; Hypsodonty based ecomorphic information of fossils of cervids….. 102 Figure 4.38; Hypsodonty based ecomorphic information of fossil suids……… 103 Figure 4.39; Hypsodonty based ecomorphic information of fossil equids………. 103 Figure 4.40; Hypsodonty based ecomorphic information of fossils of rhinoceroses……………………………………………… 104 Figure 4.41; Mean hypsodonty ranges for Siwaliks of Pakistan………. 104 Figure 4.42; Map showing estimated precipitation ranges for Siwaliks of Pakistan………………………………………………………… 105
Figure 5.1; Neogene record of CO2 and temperature change, including evidence for arctic glaciations………………………………………… 155 Figure 5.2; Neogene record of changes in vegetation structure in Siwaliks…… 155
Figure 5.3; Predicting C3/C4 dominance of grasses related to temperature
and pCO2………………………………………………………… 156 Figure 5.4; Relationship of mean hypsodonty index (HI) to diet and habitat type… 156 Figure 5.5; Mean hypsodonty based paleocommunities succession in the Siwaliks of Pakistan…………………………………… 157 Figure. 5.6; Carbon isotope record from pedogenic carbonates and mammalian dental enamel………………………………… . 157 Figure 5.7; Species richness (diversity) in ungulate remains through Siwaliks series of Pakistan……………………………………… 158 Figure 5.8; Number of murid and cricetid rodent species found or inferred to be present between 18 and 1.5 Ma in the Siwalik sediments………………………………………. 158 Figure 5.9; Relative abundance among ungulates and succession of paleocommunities during Siwalik chronology………………… 159 LIST OF APPENDICES Appendix 1; Locality database of mammalian fossils from Siwaliks of Pakistan….… 189 Appendix 2; Mesowear Type III data of living and fossil species of mammals.…. 193 ix
Appendix 3; Microwear data for Siwalik ungulates……………………………. 200 Appendix 4; Comparison of microwear data by two persons (NS magnification vs. MT magnification)……………………………………… 205 Appendix 5; Ecomorphic and biostratigraphic data of Siwalik anthracotheres and proboscidea……………………………. 209 Appendix 6; Faunal list of primates, lagomorphs and carnivores of Siwaliks…………………………………………………… 211 Appendix 7; Raw mesowear data of Giraffokery punjabiensis, Tragoportax sp., Selenoportax sp., Gazella lydekkeri……… 212 Appendix 8; Hypsodonrty measurements of boselaphines incorporated for mesowear analysis…………………………………… 214 Appendix 9; Biostratigraphic ranges of fossils species of ungulates from Pakistan Siwaliks…………………………………………… 215 Appendix 10; Mesowear database of extant species of ungulates………………… 222 Appendix 11; Microwear database of extant species of ungulates……………… 226
Appendix 12; Morphometric and ecomorphic database of living species of ungulates……………………………………………… 229
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LIST OF ABBREVIATIONS
Ma: Million years ago Myr: Million years MN: European Mammal Neogene zone scale NMU: Neogene Mammal Unit used for Chinese chronology N: Numbers AMNH: American Museum of Natural History, New York BMNH: British Museum of Natural History, London GSP: Geological Survey of Pakistan, Islamabad and Quetta GCUPC 1295/09: Government College University, Lahore Paleontological Collection, department of Zoology, Pakistan, in the text the upper figures of the serial numbers represent the collected number of the specimen and the lower ones denote the collection year. GIS: Geographic Information System. PUPC 04/11: Punjab University Palaeontological Collection, department of Zoology, Punjab University, Lahore, Pakistan, in the text the upper figures of the serial numbers denote the collection year and the lower ones denote the collected number of the specimen of the respective year. HMV: Harvard Museum of vertebrates, USA. NS: Nikos Solounias ZD: Zoology Department. M2: Second upper molar m3: Third lower molar Frmn: Formation.
xi
ABSTRACT
Ungulate remains from Siwaliks of Pakistan provide a long and continuous record of diverse geochronologic ranges and ecological niches that in turn help to explore paleoenvironments of Pakistan Siwaliks. Ecomorphic data of ungulates dental material via hypsodonty, dental microwear, mesowear type-I, II and type-III methods was collected in the present study and on comparison of dental ecomorphic data of 160 taxa of ungulate remains with standardized data of their extant communities revealed a baseline data which helped for reconstruction of paleoenvironment of the target area. In the early Miocene (18.3-15 Ma), the paleocommunities of suids, tragulids, giraffids, anthracotheres and boselaphines have been found to give rise to 2 lineages each. However, rhinos and proboscideans have evolved into 3 to 4 lineages. Ecometric analysis has shown the predominance of browsers (50%), followed by fruigivores (16.7%), mixed feeders in closed habitat (16.7%), and omnivores (15-17%) in forested habitats. In the middle Miocene (14.2-11.2 Ma), the paleocommunity has exhibited maximum species diversity that documented 54 lineages of ungulates including Sivapithecus sp. Un till 11 Ma, on the basis of baseline data the proportions of browsers gradually decreased, forest fruigivorous and mixed feeder in closed habitats increased, however, the lineages showed no significant changes in their dietary adaptations. By 11 Ma, mixed feeding in open habitat taxa appeared with predominance of forest fruigivores (35%) and browsers (32.5%). There was decreasing proportions of frugivore/selective browsers (35-16%), browsers (32-23.3%) and mixed feeders in closed habitats (19-16%) and increasing prevalence of mixed feeders (2.7-28%) in open habitats and grazers (0- 4.64%) at 8 Ma. The latest Miocene (8-6.5 Ma) of the Siwaliks chronicled the progression of great faunal turnover event during which a significant number of long lasting lineages belonging to hipparionine horses, rhinoceroses, boselaphines, sivatherines, antelopes and tragulids altered their feeding adaptations from browsing to mixed feeders in open habitats/grazers. Most of the lineages of mixed feeders in closed habitats and fruigivores gradually became extinct before 7 Ma, whereas, successive influx of lineages of mixed feeders in open habitats, grazers with stable turnover of browsers and omnivores progressed during 6.5-0.5 Ma. xii
The mesowear, microwear data from late Miocene and hypsodonty based ecomorphic data (18.3-0.5 Ma) has provided succession of the paleocommunities in Siwaliks which portray the evolution of the siwaliks ecosystem depicting the change from closed vegetation system (18.3-8.5 Ma) to semi-closed one and from semi-closed vegetation (8.5-6.5 Ma) to open vegetation system (6.5-0.5 Ma). The climate appears to have been evolved from humid and warm to dry seasonal and monsoonal one. The paleoclimate and vegetation succession has been found to lead to a cascade of diverse environmental mosaics ranging from tropical multi-canopied forest in the early Miocene to tropical evergreen forest during the middle Miocene. Thenceforth, moist deciduous canopy forest (11-10 Ma), dry deciduous forest (at 9 Ma), mosaics of dry deciduous forest and temperate woodland (at 8.5 Ma), woodlands with limited patches of deciduous forest, wooded savannas (8.5-6.5 Ma) progressed. The wooded savannas with guilds of grassy savannas (6.5-4 Ma) interspersed with deciduous forests and woodlands (4-3 Ma) and mosaics of wooded and grassy savannas (3-2 Ma) ecosystems evolved. The disappearance of most of the forested patches and prevalence of pure grasslands occurred during 2-0.5 Ma. This study provides a comprehensive account of the paleaoenvironment of Pakistan Siwaliks in relation to mammalian biostratigraphic and paleoecologic processes at an evolutionary scale.
1
CHAPTER 1 INTRODUCTION Siwalik Hills are freshwater sedimentary deposits exposed throughout the Northern areas of Pakistan, India and Southern flanks of Nepal and Bhutan (Acharyya, 1994; Nelson, 2003). In Pakistan, these deposits encompass the Potwar Plateau and Sub- Himalayan areas that are particularly well exposed in Potwar Plateau in vertical and horizontal sections (Barry et al., 1995, 2002; Nelson, 2003, 2005) (Figure 1.1). Geological history of Siwalik Hills has witnessed a diversity of climatic changes driven by tectonic movements in different directions along the meridians of longitude and across the parallels of latitude. The paleoclimate has been changing throughout the geological history of Siwalik Hills. The orogenesis and solar radiations developed a wide range of landscapes and thus a diversity of fauna and flora. Major alterations in paleoclimate have probably been the most important factor determining changes in paleoenvironment of Pakistan Siwaliks mainly revealed by evolutionary progression of fossil record of ungulate mammals. Reconstruction of paleoenvironmental evolution of Pakistan Siwaliks at temporal and spatial resolution based on ecomorphic data explored via hypsodonty, dental mesowear and microwear methods on herbivorous mammals has been addressed in this dissertation. The freshwater fluvial deposits of Potwar Siwaliks document most diverse, continuous and longest array of terrestrial and freshwater records of fossils, mainly comprising of Neogene mammalian remains, in the world (Barry et al., 2002; Patnaik, 2003; Badgley et al., 2005; Patnaik, 2011). Hence, these deposits provide an excellent laboratory for studying biochronostratigraphy, systematics, palaeoecology, palaeoclimatology and fluvial dynamics (Patnaik 2003, 2011). The lateral exposure and good chronostratigraphic control of these deposits allows detailed study of sediments and ancient biota at spatio-temporal scales which in turn offers changes in faunal community structure and habitats so significant for treating these sediments as a model to discern faunal dynamics and terrestrial paleoenvironments of Eurasia and other contemporaneous landscapes in the world (Barry et al., 2002; Badgley et al., 2005, 2008). These deposits have been focus of attention by the national and international workers in paleontological and paleoecological research for the last few decades (Barry et al., 1982; Flynn et al., 2
1990a; Barry et al., 2002). Most of them were interested to investigate how environmental change affected mammalian evolution in the Late Miocene, evolution and faunas of Miocene Siwalik sequence, and to decipher the evolutionary history of different groups of mammals with reference to taxonomic and distribution context. However, the exploration of paleoenvironment of Siwaliks of Pakistan as a whole on the basis of ecomorphic adaptations in ungulates was not investigated or investigated poorly. This region is promising for reconstruction of paleoenvironmental sequences by investigating the evolution of dietary adaptations of ungulates at spatio-temporal scales. Therefore, this novel research project on paleoenvironmental evolution of Pakistan Siwaliks needs to be worked out on the basis of ecomorphic data of extinct and extant ungulates. The geological history of the Siwalik Hills is closely linked with the cascades of tectonic events that happened due to the disappearance of Gondwanaland, Tethys Sea, uplift of the Himalayas and Tibetan Plateau (Pilgrim, 1913, 1919; Pascoe, 1920). About 100 Ma, Indian plate laid far to the South Pole, rotated away from Madagascar and began to move northwards and finally collided with Asia (ca. 25 Ma) when the great Late Oligocene and early Miocene orogenesis of Himalayas began (Irving, 1977). During the period of Siwalik deposition the drainage system of Northern India was opposite to its present configuration. A great "Siwalik River" or "Indobrahm River" used to flow northwards to the remnants of Tethys Sea. As the sea retreated into the present confines of Indian Ocean, the Siwalik River followed it turning it at right angles from its northward course, and following the present Indus Valley to the Arabian Sea. It was along this Siwalik River that the Siwalik sediments deposited (Pilgrim, 1913, 1919; Pascoe, 1920). These Siwalik Hills are foothills of Himalayas encompassing Potwar Plateau and Sub-Himalayas. The Potwar Plateau lies between the northern slopes of Salt Range and the southern side of Rawalpindi district (Figure 1.2). The geological formations of Siwaliks comprise Neogene fluvial deposits widely exposed throughout the Indo-Pak Subcontinent along the northern and western flanks paralleling the collided zone between Indian and Asian plates. The Siwalik deposits in Pakistan are well exposed on the Potwar Plateau displaying a folded belt extending from the Salt Range in the south and the Margala Hills in the north and from the Jhelum River in the east to the Indus River on the west (Barry et al., 2002) (Figure 1.2). The Salt Range forms a separate line 3
of low flat-topped hills and tending in a slight southwesterly direction, along the northern side of the Jhelum River. The fossils recorded from the Salt Range of Punjab have found to be immediate antecedents and in many instances ancestral directly to the specific vertebrate groups of Siwaliks. The level reaches of Potwar Siwaliks merge into gently dipping strata which form the summit of the Salt Range. Lithologic profiles of Siwaliks include siltstones, sandstones, clays, rare marls and mudstones (Barry et al., 2002). The Potwar Siwaliks is classified into six or more geological formations (Cheema et al., 1977) based on relative proportions of fine-grained sediments, mudstones, silt and the size of large sandstones. These lithological variations may be linked with changes in biotic structure, taxonomic composition and the mode of organic preservation (Willis and Behrensmeyer, 1995). All the geological formations seem to be time-transgressive to some extent, though it is not shown (Barry et al., 2002). Pronounced changes in major sedimentologic units have been observed within the Siwalik stratigraphic succession at spatial resolution of Kilometers to many tens of Kilometers horizontally and ten to hundreds of meters vertically (Willis and Behrensmeyer, 1995). Sedimentologic trends chronicles basin-wide changes in palaeoenvironmental conditions concerning tectonically regulated sediment accumulation rates, basin subsidence and regional climate change (Willis and Behrensmeyer, 1995). Depositional environments of Siwaliks comprises of small to large water channels, palaeosols, levees, and swamp deposits or rare ponds. Such environments are evident in all the geological formations Siwaliks but with varying proportions of occurrence. The geological formations of Pakistan Siwaliks are comprised of sediments of ancient rivers which deposited throughout the early Miocene to late Pliocene and mid Pleistocene (Barry et al., 2002). Sedimentological and taphonomic investigations reveal that the vertebrate remains of Siwaliks are documented in all lithologies, most frequently found in main channels, secondary channels, levees, in food deposits of the streams (Badgley and Behrensmeyer, 1980; Badgley, 1986a, b; Behrensmeyer, 1987) from foodplain ponds and palaeosols (Patnaik, 1995; Badgley et al., 1998) but are rarely found in the thin marls accumulated in shallow ponds as attritional assemblages mainly derived from their surroundings, however, it seems that paleo-habitat specific associations of mammalian remains have not 4
been preserved (Badgley and Behrensmeyer 1980; Badgley et al., 1995). Fossil vertebrates occur as concentrations of disarticulated, often fragmentary bones in main channels whereas in the small floodplain channels, a very few concentrations contain 1000 or more specimens, but the majority of concentrations have only 5-200 fossils. Because the large channel, levee, and paleosol facies contain only isolated specimens or small, low-density scatters, our collections of fossils from the different facies are biased toward over representation of small channel floodplain sites. This bias, in addition to those introduced by taphonomic processes common to the different facies, places strict limits on the ability to reconstruct the living communities from the fossil assemblages (Badgley, 1986 a, b) and draw meaningful conclusions from the occurrences of individual taxa. Specific concerns include uneven sampling through time, differential preservation of larger-bodied animals and durable parts such as teeth, errors in age-dating imposed by uncertainties in correlation and paleomagnetic timescale calibrations, and uneven taxonomic treatment across groups. Changes in the depositional systems of the Pakistan Siwaliks are coincided with the changes in paleoclimate as well as vegetation mosaics during the Late Miocene. Nagri Formation large emergent river systems were replaced by Dhok Pathan Formation interfan river systems beginning around 10.1 Ma (Willis and Behrensmeyer, 1995; Barry et al., 2002). After 9.0 Ma, these interfan river systems were less well drained, with more seasonally variable flow and more avulsions. Vegetation probably responded to specific local conditions, resulting in mosaics over distances of 100s of meters to a few kilometers (Barry et al., 2002). It has been found that most of the localities were deposited over intervals of tens to thousands of years, revealing that single localities may document organisms that perpetuated several thousands of years apart, probably in different paleohabitats and ecological paleocommunities (Barry and Flynn, 1990). Recent taphonomic investigation of fossils of small mammals and microvertebrates indicate that the fossils were accumulated by carnivorous mammals and birds (Patnaik, 1995; Badgley et al., 1998). A great diversity of fossils of vertebrate groups mainly includes turtles, freshwater fishes, lizards, snakes, aquatic birds, crocodiles and mammals. Mammalian remains are documented by >50,000 catalogued specimens of terrestrial habitats that dominate the Siwalik fossil record in the floodplain of Potwar Plateau (Badgley et al., 2008). Major 5
groups of mammals are comprised of commonly occuring perissodactyls, artiodactyls and rodents while uncommon groups are proboscideans, carnivores, primates, and rare creodonts, aardvarks, lagomorphs, and tree shrews (Barry et al., 2002; Badgley et al., 2005, 2008). As regards ungulates, Siwalik record of Pakistan documents several species of gomphotheres and deinotheres; more than half dozen of species of rhinoceroses, one lineage of chalicothere, and notable number of hipparionine horses; very diverse artiodactyls, comprising of > 20 species of suids, 14 lineages of tragulids, and nearly 41 lineages of bovids, 5 species of cervids, 9 lineages of giraffids. Among other goups; a substantial diversity of rodents, including hamsters, squirrels, true mice, bamboo rats, hedgehogs, rare rabbits and pangolins have also been reported. About 75% taxa of the mammalian remain have been taxonomically well resolved (Badgley et al., 2005, 2008). As a result, mammalian species richness and their faunal turnover can be investigated for all the major groups except the proboscideans, the rhinos, the carnivores, and the few of the small groups of mammals (Badgley et al., 2005, 2008). 1.1. CONSTRUCTION OF CHRONOSTRATIGRAPHIC FRAMEWORK The refined chronology of Siwalik lithology and biostratigraphy is based on collecting additional fossils and a direct consequence of extremely careful documentation of provenance information calibrated with a precise paleomagnetic framework. Chronological control is a prerequisite for studying pattern and rate of evolution within and across the mammalian lineages. 1.1.1. LITHOSTRATIGRAPHY Owing to lithostratigraphic nomenclature, the Siwalik deposits are divided into three groups 1) Lower Siwaliks 2) Middle Siwaliks 3) Upper Siwaliks (Figure 1.4). The Lower Siwalik Group consists of the Kamlial and Chinji formations. The Middle Siwalik Group consists of the Nagri and DhokPathan Formations. In the Potwar Plateau region, the Upper Siwalik Group consists of sediments that have been referred to the Soan Formation (Tatrot, and Pinjor formations) (Barry et al., 1980). Pilgrim (1926a) proposed faunal zones on the basis of which lithological units were recognized (Figure 1.4). The development of biochronology in the Neogene of Siwaliks proceeded differently from that in North America and Europe because of prevailing different set of paleoenvironmental conditions. Colbert (1935) revised the biostratigraphic correlation of 6
Pilgrim (1913) and compared it to stratigraphic equivalents in America and Europe. Magnetic polarity stratigraphy has shown that certain lithostatigraphic horizons and contacts are diachronous except in local sections. Used in a very broad and loose sense as stages, Pilgrim's faunal zones have been and still are useful although it is apparent on close scrutiny that there are fundamental problems associated with their continued use apart from the confusion resulting from lithostratigraphic and bio- or chronostratigraphic terminology. Measured marker stratigraphic horizons from the lateral exposures (of Khaur Area) have been used to establish lithostratigraphic correlations between local, regional and interregional sections. Such lithologic correlations can provide only rough estimates of relative age relationships between faunal events or facies changes in different channels (nalas or Kas as pronounced by local vernaculars) of Khaur region. Correlations to even more distant Siwalik lithofacies corresponding to geological formations like Chinji or Nagri reflect far more time differeces (Barry et al., 1980). The intention of previous workers to consider lithological sections as chrono-specific entities has posed problems nomenclature and interpretation that need to be unraveled completely (Pilbeam et al., 1979). The establishment of lithologic boundaries as reference points to explore the fossil specimens is important when these are hypothesized to be more or less isochronous. However, the magnetostratigraphic calibration of these lithological reference points reflect that the sections are very complex as well as time transgressive ranging 30-40 km area with chronologic difference on the sequence of 1.0-1.5 my (Barry et al., 1980). 1.1.2. BIOSTRATIGRAPHY Biostratigraphy of Siwaliks has been explored on the basis of faunal interval zones, observed/inferred first appearance datum (FAD) and observed/inferred last appearance datum (LAD), stratigraphic ranges of mammalian fossil taxa, occurrence of zonal marker species or faunal assemblages in a particular faunal interval zone. The appearance and disappearance of marker vertebrate taxa can be considered as firm baseline for comparison with other fossil yielding regions of Siwaliks of India and Nepal (Barry et al., 1980). Biostratigraphic framework may guide assessing the changes in mammalian community structure and patterns/episodes of faunal turnover. An understanding of litho-and chronostratigraphy may help studying the evolution of 7
terrestrial paleoenvironment and faunas during Siwalik succession (Barry et al., 1980). The significant changes in biostratigraphic record of the Siwaliks correspond to dynamic changes in major global climatic, tectonic and oceanographic events. Appearance refers to speciation and immigration events. Disappearance signifies regional and global extinction events. In this biostratigraphic analysis, about 205 mammalian fossil taxa belonging to eight (08) orders; Insectivora, Primates, Perissodactyla, Artiodactyla, Lagomorpha, Rodentia, Carnivora, Elephantoidea, and Rhinocerotidae have been included (Appendix 5). For each taxon, first or last occurrence (FO) or (LO) event has been defined. Gomphotherium s. l. Interval Zone (20 Ma-16.3 Ma); This interval (20 Ma-16.3Ma) is designated by African immigrants Proboscidean taxa i.e. Gomphotherium and Deinotherium as zonal marker elements together with remains of rhinoceroses, carnivores, suids; Paleochoerus pascoae, Palaeochoerus perimense, Bunolistriodon sp., early bovid; Eotragus noyei, and some gigantic anthracotheres (Sarwar, 1974; Solounias et al.,1995; Ahmad, 1995). These mammals were small to medium in body size and showed browsers to frugivorous dietary adaptations. Listriodon sp. Interval Zone (16.3 Ma-14 Ma); The fauna recovered from this interval (Chinji Formation) mainly consists of Listriodon sp. of suids, and taxa of ruminants, rhinoceroses, rodents, primates and tragulids. The faunal list comprises of thyronomyid, rhizomyid, ctenodactylid, and advanced forms of muroid rodents along with species of Listriodon, Brachypotherium, Dorcabune, Sanitherium, a large giraffoid and gibbon like hominoid. This faunal list may be compared with the other middle Miocene Siwalik localities. Listriodon pentapotamie ranges from Upper Kamlial to Late Chinji in stratigraphic configuration. Nearly all of the ungulates were closed habitat taxa with browsing, forest frugivorous and browsing mixed feeders adaptations. Giraffokeryx punjabiensis Interval Zone (14.0 Ma-9.5 Ma); The Giraffokerycinae are documented sporadically in the Chinji Formation of the Lower Siwaliks (Colbert, 1935; Bhatti et al., 2007). Giraffokeryx punjabiensis has already been mentioned several localities of the late Middle Miocene age (Bhatti, 2005), occupying a wide territory from Western Europe to India (Bohlin, 1926; Bosscha-Erdbrink, 1977; Gentry et al., 1999; NOW database 2013). The Deinotherium pentapotamiae and Listriodon pentapotamiae 8
are characteristic markers of the Chinji Formation (14–11.2 Ma). Helicoportax is also a representative of the upper Chinji Formation (Pilgrim, 1937, 1939). The long chronologic and wide geographic ranges of Listriodon pentapotamiae together with Giraffokeryx punjabiensis, Gaindatherium browni, Brachypotherium fatehjangense, Deinotherium pentapotamiae, and Dorcatherium sp. reveals a middle Miocene age of the Chinji Formation. The fauna is in favor of a late Middle Miocene age because the comparison of the material with several representatives of the fauna indicates a middle Miocene age (Pilgrim, 1937, 1939; Heissig, 1972; Thomas, 1984; Pickford, 1988). The ungulate taxa were predominantly forest frugivores/selective browsers, browsers and browsing mixed feeders. Hipparion s. l. Interval Zone (9.5 Ma-7.4 Ma); It is the most significant event of faunal turnover and is defined by North American immigrants of hipparionine horses together with the appearance of new forms of suids and giraffids from Eurasia as well as rodents and tragulids from Africa. This interval also marks the extinction of local fauna of large herbivorous mammalian fossil taxa such as Listriodon, Conohyus, Giraffokeryx, numerous bovids and tragulid species. It should be noted that the known ranges of hipparionine horses in deposits of the Potwar Plateau (Hipparion s.l. of Barry et al., 1982) is from 9.5 Ma (FAD or "first appearance datum") to 1.5 Ma (LAD or "last appearance datum"). The Hipparion s.l. Interval Zone (from 9.5 to 7.4 Ma) is a subset of the observed range of hipparionine horses in the Potwar Plateau succession. The base of this zone is defined as the FAD of Hipparion s. l. (Barry et al., 1982), and the top of this zone terminates just below the FAD of Selenoportax lydekkeri. This interval chronicles the increasing progression of grazers, open habitat mixed feeders with diminishing of browsing mixed feeders, frugivores and browsers. Selenoportax lydekkeri Interval Zone (7.4 Ma-5.3 Ma); A major faunal turnover starting at 7.5 Ma changed the taxonomic composition of mammalian fauna of the Siwaliks, making it much similar to the contemporaneous faunal elements of western and northern Eurasia (Barry et al., 1991). During this last mammalian interval of Miocene, the faunas exhibit characteristics of more woodland-open habitat taxa belonging to very diverse families of Hyanidae, Felidae, Equidae, Rhinocerotidae, Bovidae, Suidae and Giraffidae (Khan et al., 2011a). The biostratigraphic range of Selenoportax in the 9
Siwaliks is from the Late Middle Miocene to the Pliocene (Akhtar, 1992; Khan et al., 2009a). Nevertheless, Barry et al., (2002) recognized a maximum range of 10.3 to 7.9 Ma for this taxon. Qiu and Qiu (1995) list Selenoportax sp. from the Lufeng fauna (Chinese Miocene), age of 11.1-8.0 Ma (Flynn and Qi, 1982; Steininger, 1999). Pachyportax, on the other hand, is considered as a typical Late Miocene taxon. Pachyportax occurs in the Nagri and the DhokPathan zones of the Siwaliks, until Proamphibos replaces it soon after the start of the Tatrot. Hence, a Latest Miocene date around 7.0 Ma (Barry et al., 1991) would be considered as a possible date for a fauna containing P. latidens. Overall, the DhokPathan Formation faunas resemble those from the Turolian Land Mammal ‘Age’ as defined in Europe, North Africa and West Asia. There rather restricted time range seems to fall close to the boundary between the Vallesian and the Turolian Land Mammal ‘Ages’. Therefore, the age of the DhokPathan, is considered as the Late Miocene. Barry et al., (2002) suggested that the age of the DhokPathan was the Late Miocene. However, the cervids, the bovids, the giraffids, and the suids suggest a Late Miocene to early Pliocene age (Khan et al., 2009a, b, c, 2011a, b). The faunal association of the DhokPathn stratotype indicates an age to date between the Late Miocene and Early Pliocene (Figure 1.4). This interval exhibits the prevalence of hypsodont and open habitat mesodont ungulates and gradual disappearance of selective browsers and browsers. The Upper Siwaliks fluvial sequence of the Indo-Pak Sub-continent is one of the most continuous of its age, spanning in time from the Late Pliocene up to the Middle Pleistocene, ca. 3.3-0.6 Ma (Behrensmeyer and Barry, 2005; Dennell et al., 2006, 2008; Nanda, 2008). In the local lithostratigraphy the Upper Siwaliks comprises from the base to the top of the Tatrot Formation, the Pinjor Formation and the Boulder Conglomerates (Nanda, 2002; Kumaravel et al., 2005; Dennell et al., 2006). Hexaprotodon sivalensis Interval Zone (5.3 Ma-2.9 Ma); Fossil material attributed to Hexaprotodon sivalensis, Proamphibos lacrymans, Stegodon sp., Potamochoerus sp., Dorcatherium sp., Hippopotamodon sivalensis, Percrocutta grandis, Bison cf. sivalensis, Proamphibos sp. “Rhinoceros” sivalensis, Tetralophodon falconeri and Anancus sivalensis has also been recovered from the same deposits (Sarwar 1977; Akhtar 1992). The lower boundary of the Tatrot Formation ranges between 3.5-3.3 or 3.4-3.2 M. y (Hussain et al., 1992; Barry et al. 2002) that corresponds to the lower part of the Gauss 10
magnetochron, whereas the upper boundary of the Tatrot Formation between 2.4-2.6 My (Kumaravel et al. 2005; Dennell et al., 2008; Nanda, 2008). Hence, Tatrot deposits are roughly correspond to the latest Pliocene. The reffered taxa recovered from deposits in the vicinity of Tatrot and Kotal Kund villages belonging to the Tatrot Formation, while sequences of the younger Pinjor Formation are exposed more to the south (Shah, 1980; Johnson et al., 1982; Dennell et al., 2006). The fossiliferous deposits of the Tatrot Formation outcropping in the area consist of pale pinkish-orange brown clays, brownish grey siltstones and shale, and greenish grey fine to medium grained sandstones intercalated with dark grey conglomerates. The faunal composition shows the dominance of open habitat taxa and rare occurrence of frugivores and browsers. Elephas planifrons Interval zone (2.9 Ma-1.5 Ma); The Pabbi Hills are probably representative of Pinjor deposits of Upper Siwalik in Northern Pakistan which are endowed with Elephas planifrons as regional marker chrono-taxon accompanied by Elephas hysudricus, Equus sivalensis, Hypohyus sp., Sivatherium giganteum, and cervids. Numerous taxa of Artiodactyles went extinct upon the first appearance of proboscideans and suids from Africa and cervids and equids from Eurasia. The modern faunal elements of Indo-Pak sub-continent made their origin from this last episode of faunal turnover. Most of the Siwalik ungulates inhabiting in this interval had adapted to open habitat niches displaying mixed feeding to grazing life style. This study incorporates the synergistic ecological interplay of extinct and extant communities of ungulates as baseline for exploration of paleoenvironment of Pakistani Siwaliks. The rational for selection of ungulates is that they are abundant, have extensive geographic and chronologic ranges, morphologically diverse and have been adapted to a broad range of habitats and feeding strategies. These attributes can capture diverse environmental variations in temporal and spatial scales. This study has been focused on dental material of ungulate remains from Siwaliks of Pakistan to explore their diet and paleoenvironment as a whole due to their ubiquity and close relationship to understandable physical and biological interactions. The dentition can serves as interface between an individual and his environment as the orientation of the molar teeth reflects taxonomic signals and adaptations to the dietary niches. The investigation of the dental variation can provide insight into the eco-morphological space for sympatric lineages of 11
ungulates. Whatever an organism eats and drinks ends up in its teeth and bones. The teeth of ungulates grow for several years creating an environmental record along their length, a "tape recorder" of the changing environments or seasons. Thus, fossil teeth are useful recorders of the paleoenvironment in which the animal thrived and demised. The simple expressions of bio-mechanically induced tooth wear patterns were proved to have strong potentialities for resolving the spectra of dietary inferences among ungulates (Fortelius and Solounias 2000). The ungulate remains are the best indicator of great diversity of abiotic and biotic variables as the details of regional ecology is recorded in their dental morphology (Fortelius et al., 2002). The mean value of hypsodonty in herbivore community is indicative of the precipitation level of local habitat of that community (Damuth and Fortelius, 2001). The application of different research tools such as hypsodonty, microwear and mesowear on dentition of extinct ungulates from Siwaliks has provided useful information regarding their paleodiets and paleoecological interpretations. Paleoecology of 163 taxa of ungulates referred from 97 fossil localities has been explored by considering hypsodonty based ecomorphic information (diet, habitat etc.) The mesowear study of 17 ungulate species from Miocene (18-5 Ma) and microwear of 12 species from Late Miocene of Pakistan have also been worked out for their paleoecological reconstructions. Hypsodonty is recognized as an adaptation for grazing and has been used as morphological proxy to interpret habitat selection and feeding preferences in herbivorous mammals (e.g. Janis et al., 2000, 2002; Feranec, 2002; Jernvall and Fortelius, 2002; Feranec, 2003; Fortelius et al., 2003; Hopkins, 2003; Palmqvist et al., 2003; Bargo et al., 2006; Stromberg, 2006). Hypsodonty may be referred as multivariate ecological proxy for diet, habitat, vegetation, climatic regimes in herbivorous fossil mammals for unraveling environmental reconstructions (Fortelius et al., 2002, 2003). The functional interpretation of hypsodonty may explore some classic examples of evolutionary change (e.g. the diversification of equids, giraffids and the development of the grassland paleoecosystem in Pakistan Siwaliks and in its contemporaneous landscapes through middle Miocene to Pleistocene times (Simpson, 1951; Shotwell, 1961). It has been hypothesized that hypsodonty evolved as a result of co-evolution of phytolith, grit, dirt laden open vegetation and herbivory. 12
The widespread evolution of hypsodonty among Siwalik ungulates appears to be coincided with the cascade of events of floral progressions from closed habitats to open habitats during early Miocene to Pleistocene. It was relatively constant in succeeding genera of early to mid Miocene but there was a more significant acceleration in Late Miocene to Pleistocene. Degrees of hypsodonty include (1) brachydont (low crowned teeth) (2) mesodont (middle crowned teeth), and (3) hypsodont (high crowned teeth) (MacFadden, 1998; Fortelius et al., 2002; Damuth and Janis, 2011). Brachydonts correspond to browsers whereas mesodonts and hypsodonts signify mixed feeders and grazers respectively. Browsers indicate forested habitat, mixed feeders represent forest, woodlands to savannas while grazers show savannas to grasslands (Fortelius et al., 2002, 2003, 2006; Damuth and Janis, 2011). The three major dietary categories have been considered as a model for interpretation of paleo-habitats and paleodiets leading to reconstruction of paleoenvironmental conditions (e.g., Janis, 1988; Quade et al., 1992; Solounias et al., 1995; MacFadden and Cerling 1996; Janis et al., 2000). The hypsodonty of ungulates has been used to broadly map paleoenvironmental changes through Siwalik succession. The inference of paleohabitats or regional paleoenvironment is corroborated by correlating the mean hypsodonty values with the patterns of hypsodonty values and habitats of extant taxa (Fortelius et al., 2002, 2003, 2006). The major problem for examining the resolution of paleodiet/paleohabitat categories of the fossil species of ungulates in the Siwaliks of Pakistan is that its detailed record of Neogene paleovegetational changes is insufficient (Jacobs et al., 1999; Stromberg, 2002, 2004). The reason why this study addresses the correlation between htypsodonty based dietary/ habitat categories in extant ungulates to validate the paleoecological inferences of related extinct taxa (e.g. Janis, 1988, 1995). Hofmann and Stewart (1972) suggested further subcategories within the three main categories to reflect some of this complexity. Thus, concentrate selectors can be subdivided into tree and shrub foliage eaters and fruit and dicot foliage eaters. Grazers include the subcategories fresh grass grazers (FG), and dry region grazers (GG). The mixed feeders comprise the subcategories preferring grasses= mixed feeders in open habitats (MFO) and preferring forbs, shrubs, and tree foliage= mixed feeders in closed habitats (MFC). These subcategories have been used in paleecological studies to 13
determine the percent of a particular forage type within the diet of an extinct ungulate in the Siwaliks. Previous studies have shown that common fossil species of mammals (common tooth crown types) drive the evolutionary increase in hypsodonty (Jernvall and Fortelius, 2002, 2004). Hypsodonty in many ungulate lineages shows a strong relationship with local mean annual precipitation in extant mammalian communities (Damuth and Fortelius, 2001). Mean Hypsodonty patterns explore the conditions of the vegetation which may be nominated as “generalized water stress” depicting the humidity-aridity gradients. Mean Hypsodonty has direct proportion with arid conditions and inverse proportion with humid environment. This implies that the higher value of mean hypsodonty is indicative of more arid conditions, whereas low hypsodonty value represents more humid environment (see Fortelius et al., 2002, 2006 and Eronen, 2006 for further description). The investigation was constrained to localities of Pakistan Siwaliks, and only large mammals were selected and classified as plant-eaters or plant-dominated omnivores (suids) for analysis. The decrease in hypsodonty (brachydonty), mean body size and shift in dietary preferences
(browsers/ closed vegetation/ C3 vegetation) implies climatic warming, precipitation increase, probably spread of forests. Hypsodonty increased with altitude, which suggests
change in dietary regimes from browsers to grazers (open and dry habitats/ C4 vegetation), climatic cooling, decrease in precipitation and emergence of grassland ecosystems. The dental microwear technique is a dietary analysis that quantifies the microscopic wear on teeth of herbivorous mammals. The development of microscopic scratches and pits on the tooth enamel during the feeding process of herbivores is called microwear. This study incorporates microwear of 12 fossil species of ungulates from Late Miocene of Pakistan to extrapolate their paleodiets. Studies on microwear of extant herbivores with known diets have revealed a correlation between microwear and diet (Teadford and Walker, 1984; Grine, 1986; Teaford, 1988; Solounias and Hayek, 1993). One noticeable trend is that grazers typically bear more scratches than browsers while browsers have more pits than grazers (Solounias and Semprebon, 2002). These observations have led to the establishment of a Microwear Index (MI) (MacFadden et al., 1999b). For a stated area of enamel (e. g., 0.5 mm x 0.5 mm), the MI is calculated as the 14
average number of scratches divided by average number of pits. As a standard, an MI below 1.5 indicates a browsing diet and an MI above 1.5 indicates a grazing diet (MacFadden et al., 1999b). Additional conclusions have been drawn by comparing the analysed tooth to a database of the microwear of extant animals with observed known diets. Bivariate plots of the average number of pits versus the average number of scratches for these extant herbivores reveal distinct morphospaces indicative of browsing, grazing, and mixed diets (Solounias and Semprebon, 2002). Analyzing microwear at only 35X magnification has been shown to reveal the same results as high magnification (Solounias and Semprebon, 2002). Solounias and Semprebon (2002) also created four additional quantifying characters to attain a more detailed understanding of dietary habits. These characters can be used in hierarchical cluster analyses to determine which kind of extant herbivores most closely resemble the fossil taxa in terms of microwear, and therefore diet. However, microwear is subject to the “Last Supper Effect,” the limitation of reflecting an individual’s last few meals, rather than the life history of diet (Grine, 1986). Therefore, microwear analyses are based on the assumption that the animal’s typical diet is reflected by these final meals. Tooth mesowear has been a useful technique in deciphering the paleodiets of various ungulates (Fortelius and Solounias 2000; Mihlbachler et al., 2011; Solounias et al., 2010; 2012). This research tool has been applied to 17 species of ungulate remains from Miocene of Potwar Siwaliks. It is based on scoring of the apices of molars from the buccal view. Tall and sharp apices are most common with browsers whereas shortened and rounded apices are more common with grazers. Two scoring scales have been developed on extant species for the evaluation of mesowear. Originally Fortelius and Solounias (2000) developed the method they treated the wear down of cusps as a two variable feature. The cusps were high or low. The actual apex was either sharp or rounded and in extreme wear blunt. Kaiser et al. (2000) and Kaiser and Solounias (2003) extended these observations to more than one cusp per individual and tested its predictability by involving more observers on the same teeth. This is mesowear I. Mihlbachler et al. (2011) used a new scoring scale where the height of the cusp was integrated with the wear of the apex into a single scale; single variable. In all these studies the paracone or the metacone of M2 is preferably used. A selection of the sharper 15
of the two is the ideal cusp. If a cusp is broken or problematic an adjacent cusp is used in the scoring. This is termed as mesowear II. There are several studies utilizing these methods (Clauss et al., 2007; DeMiguel et al., 2008; Eronen et al., 2009a, b). Throughout these studies, thousands of teeth were examined, additional significant mesowear differences were noted, which have previously not been addressed. For example, regions other than the buccal apex contained information that appeared to be significant. In a ruminant or equid tooth, four primary bands of enamel are present. The buccal band is presently termed band 1. This band was used in mesowear I and II. Going lingually, the next is band 2, followed by band 3. The innermost band is band 4. Bands 1 and 2 constitute the paracone and the metacone. These two enamel bands enclose a region of dentine. Observations (by Nikos Solounias) for many years revealed that band 2 contained better mesowear information than band 1. Band 2 shows mesowear with interesting variations; variations which could be used in differentiating in diet than those of band 1. In addition, the occlusal view of the anterior part of band 2 gave different mesowear from the posterior. Two experiments using domestic goats were devised to see if the observed patterns from the wild were reflected in an experimental setting. Goats were fed selected types of browsing and grazing plants at different time intervals, and the teeth were examined and compared with the wild ruminants. Experiments with mesowear are rarely done, and my collaboration with the team of scientists from New York Institute of Technology, College of Osteopathic Medicine, USA allowed me to investigate the dietary implications of mesowear II and III. The recent Siwalik literature offers an opportunity to evaluate faunal changes of large herbivores on a more restricted temporal and geographic scale. It has been considered as a relative measure for describing community structure/fossil biodiversity. 1.2. OVERVIEW OF PALAEOENVIRONMENT Previous interpretations of palaeoenvironment of Siwaliks has been based on isotopic studies of nature of the palaeosols (Quade et al., 1989; Cerling et al., 1993; Willis, 1993b; Zaleha, 1994; Quade et al., 1995), marine microfossils (Wright and Miller, 1993), plant material (Sahni and Mitra, 1980), and paleoclimate modelling (Kutzbach et al., 1989; Ruddiman, et al., 1989; Iacobellis and Somerville, 1991a,b; Prell and Kutzbach, 1992; Raymo and Ruddiman, 1992). The mature palaeosol horizons in 16
different geological formations in the Chinji area (15-8 Ma) have relatively constant thickness which implies the constant mean annual rainfall (Willis, 1993b). The evidences for significant paleoenvironmental change during Late Miocene come from both the stable isotopic content of fossils and paleosols (Quade et al., 1989, 1992; Morgan et al., 1994; Stern et al., 1994; Quade and Cerling, 1995) and the sediments (Retallack, 1991; Willis, 1993a, b; Willis and Behremsmeyer, 1994, 1995; Zaleha, 1997a, b). Since the pollen remains have not been recorded from Pakistan Siwaliks, wood fossils recovered from Indian Siwaliks reveal the flourishing of tropical evergreen forests together with rare perpetuation of taxa of moist deciduous forests in this region in Middle Miocene (Prasad, 1993). The most probable bio-stratigraphic range of these forests lies between 11-7 Ma (Nelson, 2005). Moist deciduous forest taxa dominated later on and then dry deciduous forest emerge (Nelson, 2003). Leaf and pollen remains explored from Nepal Siwaliks is indicative of dominance of evergreen forests followed by semi-deciduous and dry deciduous ones in vegetation during Middle to early late Miocene spanning from 11- 9 Ma (Quade et al., 1995). Corvinus and Rimal (2001) gave detailed floral record of Siwaliks of Nepal. The Bankas Formation of Nepal corresponding to Chinji Formation of Pakistan (14.2-11.2 Ma) (Barry et al., 2002) contains leaves remains of tropical evergreen forest elements recorded today in S. E. Asia only, including Dipterocarpus (Nelson, 2005). Moist deciduous taxa appeared for the first time in Lower Nagri Formation of Pakistan at 11.0 Ma that corresponds to lower Chor Khola of Nepal. Palaeoenvironment of Siwaliks of Pakistan was like a mosaic of C3 and C4 plants during Late Miocene;
however percentage of C4 vegetation (grasses) was increasing after 8 Ma (Nelson, 2005). Changes in paleovegetation mosaic were concordant with the changes in depositional system and paleoclimate throughout late Miocene (Nelson, 2005). The large emergent river systems of Nagri Formation were replaced by interfan river systems of Dhokpathan Formation initiating around 10.1 Ma (Barry et al., 2002; Willis and Behrensmeyer, 1995). Post-9-Ma “inter-fan river systems” were less drained showing more variable flow and avulsions seasonally. Progression of paleovegetation synchronized in accordance with local conditions that resulted in mosaics ranging from hundreds of meters to few kilometers (Barry et al., 2002). A shift in δ18O ratios in palaeosol carbonates began at 9.15 Ma and with samples was becoming more and more enriched 17
through times (Quade et al., 1989; Quade and Cerling, 1995). This shift in δ 18O values has been inferred as decline in rainfall but increase in its seasonality. A documentary record of foraminiferal assemblages reflects commencing of strong summer upwelling of Arabian Sea, with strengthening continued until 8.2 Ma, revealing the major intensification of “South Asian Monsoon System” (Kroon et al., 1991, calibrated to Cande and Kent, 1995). The intensification of monsoon system may elucidate oxygen shifts in Siwaliks of Pakistan, causing increased aridification, seasonality and summer rainfall that would then trigger the progression of more open habitats with C4 vegetation
at the expanse of closed habitats and C3 plants (Nelson, 2005). The changes in isotope ratios of stable carbon and oxygen (in both δ13C and δ18O values) in the Late Miocene, resulted from long term climate and vegetation changes which led to drier, more seasonal paleoclimate and extensive grassland ecosystems (Quade et al., 1989, 1992, 1995). The oxygen isotopes signatures represent significant change in precipitation patterns initiating at 9.2 Ma that might have been changed to more drier and seasonal paleoclimate. Morgan et al. (1994) has proposed that the commencing of the vegetation change at 9.4 Ma is concordant with major faunal changes, depicting the ecosystem with more grazing taxa and less woodland inhabiting fauna
(Barry, et al., 1985) and cooler and arid paleoclimate. During 8.5-6.0 Ma, C4 savannas succeeded/replaced the C3 forests and woodlands (Bagdely et al., 2008) as most of the forest frugivores and browsing lineages maintained their paleodietary habits and demised.
Other lineages who altered their paleodietary adaptations by incorporating C4 plants persisted for more than 1Ma whiles vegetation transition (Bagdely et al., 2008). Post-7
Ma plant communities comprising of predominant C3 species (closed vegetation) were
greatly demised and those having predominant C4 vegetation (open vegetation) with
grassy woodlands, originated at 7.4 Ma, while pure C4 grasses were evident at 6.9 Ma.
The shift in carbon isotopes reflected a change from C3 dominated taxa to C4 dominated communities i.e. from closed woodland and canopy forest ecosystems (closed vegetation) to more open vegetations (Quade et al., 1989; Quade and Cerling, 1995). A few paleosols may have formed under water logged, grassy woodlands, but most formed under drier conditions and closed vegetation (Cerling et al., 1997; Barry et al., 2002). The stable carbon isotope record reveals that there was paleovegetational progression from 18
origination of significant amounts of C4 grasses to floodplain habitats with extensive C4 grasslands within the interval of 8.1 to 6.8 Ma. Tropical evergreen taxa became very rare and were replaced by majority of the taxa of moist deciduous forest and first appearance of dry deciduous genera in Surai Khola which corresponds to the later Late Miocene (second half of Dhokpathan Formation) of Pakistan at ca. 7 Ma (Nelson, 2003). The notable change in paleovegetation occurred from woodlands and shrublands to svannas during 7-4 Ma as revealed by isotopic signatures of palaeosol carbonates (Quade et al., 1989; Cerling et al., 1993; Quade et al., 1995). The new interpretations concerning palaeoenvironment of Pakistan Siwaliks have been explored on the basis of evidences drawn from hypsodonty, dental mesowear, and microwear analyses of extinct and living ungulates and their faunal turnover patterns and change in community structure. The integrated analysis of these evidences suggest that there was a gradual palaeoclimatic shift during the deposition of the Siwaliks from warm, humid, and tropical (19-14 Ma) to cold, arid and sub-tropical (11-7 Ma) to even more seasonal, arid and monsoonal (7-1.5 Ma). Early Miocene of Pakistani siwaliks was predominated by browsers, forest fruigivores, omnivores and a few closed habitat mixed feeders which indicates the existence of tropical dense canopy forest. Middle Miocene (Chinji Formation) shows maximum species diversity of browsers (BB, HB), frugivores, and mixed feeders in closed habitats which are indicative of tropical evergreen forest during 14-12 Ma. There was prevalence of subtropical forested taxa at 11 Ma to moist deciduous forested ones at 10 Ma to dry deciduous forested lineages at 9 Ma followed by mosaics of moist and dry deciduous forests till 8 Ma. During 8-7 Ma interval, the significant number of the closed habitat taxa went extinct and open habitat taxa began to replace gradually which indicated that forests and woodlands were replaced by wooded savannas. After 7 Ma, browsing mixed feeding and frugivorous taxa greatly demised whereas open habitat mixed feeders and grazers successively increased. This change in trend of ecological communities indicates the intensification of monsoonal paleoclimate and a change from semi-closed vegetation to an open vegetation system (Table 5.3;
Figure 5.9). During 8.5-6.0 Ma, C4 savannas succeeded/replaced the C3 forests and woodlands (Bagdely et al., 2008) as most of the forest frugivores and browsing lineages maintained their paleodietary habits and demised. Other lineages who altered their paleo- 19
dietary adaptations by incorporating C4 plants persisted for more than 1 Ma whiles vegetation transition (Bagdely et al., 2008). The faunal interval during 5.5-4.5 Ma shows the further extinction of selective browsers/ frugivorous, browsing-mixed feeding lineages of boselaphines, rhinoceroses and appearance of open habitat taxa of cervids and boselaphines, modern monsoonal favoring grazing lineages of bovines and reduncines with persistence of dry grass grazing taxa of hipparionines. The relative abundance of browsers, frugivores, mixed feeders in open habitats, dry grass grazers and fresh grass grazers reveals the prevalence of open wooded and grassy savannas with rare patches of dry deciduous and moist deciduous forests (Table 5.3; Figure 5.9). The faunal turnover during 4.5-3.5 Ma exhibits disappearance of hipprionine horses, Hydaspitherium-Bramatherium lineage, appearance of Equus sivalensis, Equus sp., one lineage of antilopes, persistence of cervids, 3 taxa of reduncini and boselaphini, 2 taxa of each of antilopini and bovini, one lineage of hippotragini, 4-3 proboscideans, and an anthracothere. The reconstruction of ecological composition based on presence and relative abundance of mammalian remains such as ever decreasing browsers, frugivores, and increasing of mixed feeders in open habitats, dry grass grazers, fresh grass grazers depict the prevalence of grassy savannas and grasslands with rare patches of dry deciduous and moist deciduous forests (Tables 5.1, 5.2; Figure 5.9). The significant mammalian faunal turnover between (mid Pliocene- Pleistocene 3.5-2.5) the Tatrot and Pinjor (Pabbi Hills) stratotypes reveals the reduction (from three to two) in the biostratigraphic range of proboscidea, and more hypsodont grazer E. hysudricus (and S. cf. insignis) replaced E. planifrons, whereas Equus sivalensis, Equus sp. replaced Hipparion. There was appearance of D. palaeindicus and Hemibos triqueticornis (hypsodont bovids), Rhinoceros sivalensis, R. sondaicus, Sivatherium giganteum, and very common cervids, reduncini, bovini and hippotragini. The reconstruction of habitat on the basis of presence of the mammalian taxa and their abundance revealed substantial changes whiles mid Pliocene to Pleistocene of Siwaliks in virtually disappearance of forest, widening of grassy savannas and grasslands with shrinking of wooded savannas. Faunal turnover pattern during 2.5 to 0.5 Ma reflects the prevalence of mixed feeders in open habitats and grazers with small proportions of browsers and omnivores and complete extinction of fruigivores. Such ecological 20
communities suggest the prevalence of grasslands with grassy savannas and disappearance of most of the forests. RESEARCH OBJECTIVES The purpose of this research is to collect ecomorphic data from dental materials of fossil and living herbivorous mammals for exploration of paleoenvironment of Pakistan Siwaliks by framing following objectives. 1. To compare the paleodietary adaptations using quantitative dental microwear, mesowear, and hypsodonty techniques for analyzing morphological changes in mammals and examining dietary changes during significant climate events. Thenceforth, integrate the paleoecological data, and phylogenetic information to address questions of evolutionary paleoecology. 2. To reconstruct the well resolved chronostratigraphic framework based on previously published information regarding biostratigraphic and magnetostratigraphic cross-correlation of different composit sections. It will provide information concerning faunal turnover patterns, environmental change suitable for making paleoclimatic and tectonic inferences. 3. To integrate the faunal and palaeoenvironmental data in geographical and temporal context, and there after for reconstructing paleobiogeographic maps. 1.3. SIGNIFICANCE OF WORK The explored ecomorphic data will contribute to the evolution of terrestrial ecosystem database and will attract interest among paleoecologists, vertebrate paleontologists, conservation biologists, sedimentologists, taphonomists, anthropologists, archeologists and naturalists. It will add new information in the field of paleoecology. The construction of biochronostratigraphic framework may lead to a better understanding of changes or stability in paleocommunity structure, the evolution, migration and extinction of the Siwalik fauna. It may serve as a model for constructing biochronostratigraphic framework of its contemporaneous deposits and their associated evolution of community structure. The documentation of Sivapithecus sp. together with the paleoecological interpretation of its contemporaneous ungulate remains presents an interesting example for understanding the hominoid evolution. 21
A stable biostratigraphy that is well-dated and densely sampled opens many lines of research. On the level of mammalian assemblages of Potwar Siwaliks, peaks in faunal turnover vs. intervals of relative stability can be identified and dated. This historical record of faunal dynamics is relevant for the Indo-Pak subcontinent and can be related to other well dated records. On the level of lineages, historical records of individual taxa (species or larger monophyletic groups) and evolutionary patterns have been investigated and compared. It provides first detailed evolutionary study of dietary based paleovegetation and paleoclimatic regimes. Hypsodonty, mesowear and microwear analyses based dietary evolution of ungulates provides a first sole evidence for strenghtening of paleoprecipitation, paleoseasonality gradients. This study also elaborates as to how South Asian monsoon system originated at ca. 12 Ma and successively intensified during Late Miocene in the wake of tectonic uplift of the Himalayas. This endeavor presents when modern monsoon system initiated and thereafter intensified through Plio-Pleistocene and as to how this system is concordant with the assumed spread of more open habitats due to the increased tectonic upheaval of Tibetan Plateau. Newly incorporated methods of mesowear and microwear analyses provide a more comprehensive dietary interpretation of fossil ungulates than previously known. Mesowear, microwear data and their integration with previously published isotopic data (including equid intra-tooth isotopic sampling) chronicle the dating of when browsing taxa first incorporated grass into their food and when patches of C4 grasses (grasslands)
first became more prevalent. This study provides the oldest record of C4 graze explored
so far. Hypotheses concerning faunal turnover patterns as to how and why C3 plants,
transition of C3 and C4 plants, C4 grasses together with their inhabiting taxa attained dominance during different time intervals of Siwaliks have been proposed and discussed.
22
FIGURES
Figure 1.1; Map showing geographic placement of Siwalik Hills (Source; Chauhan, 2003).
Figure 1.2; Map of Potwar Siwaliks displaying major geological formations. Rectangles A, B and C represent major/type localities from where samples mammalian taxa were reffered for this study (source; Pilbeam et al., 1979).
23
Figure 1.3; Locality map of Pabbi Hills (Pinjore Formation), district Gujrat, Pakistan (source; Dennel et al., 2008). 24
Figure 1.4; Biochronostratigraphy of Siwaliks of Pakistan and its corrleation with European and Chinese chronology. Abbreviations: BIZ=biostratigraphic interval zone, NMU=Neogene Mammal Unit, MN= European Mammal Neogene zone scale. The following references were used for this compilation: Geomagnetic Polarity Time Scale, Mankinen and Dalrymple (1979); Cenozoic Epoch boundaries, Berggren and Van Courvering (1974); Lithostratigraphy, Barry et al. (1982); faunal zones, Pilgrim (1934); Interval-Zones, Barry et al. (1982); known range of all Siwalik hipparions, Keller et al. (1977) and Johnson et al. (1982); European mammal stages, Fahlbusch (1976). 25
CHAPTER 2
REVIEW OF LITERATURE
Earlier scientific research on Siwalik Hills of Pakistan provides information about its geology, paleontology and paleoecology in chronological order. The history of the Siwalik faunas witnesses turnover events, but differ in intensity, diversity of groups affected, and coincidence of first and last occurrences, probably showing different causes. The first documentary report on mammalian fossils in Siwaliks began in the mid1830’s with the published discoveries of British colonial officers (e.g. Cautley and Falconer, 1835; Baker and Durand, 1836) who primarily focused on the description of taxa. Falconer and Cautley (1846) were the pioneers for investigating geology and paleontology of Siwaliks. Pilgrim (1910, 1913) was the first to explore the biostratigraphy of Siwaliks on the basis of mammalian faunas. He divided the strata of Siwalik group into three sub-groups i.e. Lower Siwaliks (Kamlial and Chinji formations), Middle Siwaliks (Nagri and Dhok Pathan formations), and Upper Siwaliks (Tatrot, Pinjor and Boulder Conglomerates) and correlated the seven geological formations with the standard formations/stages of European Neogene. While not strictly biostratigraphic, there was a lithological component to the zonation of the Siwaliks presented by Pilgrim (1910, 1913) (Figure 1.4). Colbert (1935) revised the biostratigraphic correlation of Pilgrim (1913) and compared it to stratigraphic equivalents in America and Europe. Having presented the zonations of mammalian fossils in the Siwaliks, Pilgrim (1910, 1926a) and then Colbert (1935) explored some of their large scaled evolutionary patterns, however, they too concerned to biogeographic and phylogenetic patterns in principle. A well documentary record of Ramapithecus from Siwalik group was re-evaluated as human ancestor by Simons (1961). Many teams of international workers re-excavated the mammalian fossil fauna of Siwalik Hills during 1950-1970 (Dhem et al., 1958a, b; Pilbeam et al., 1977). There were no significant modifications of Pilgrim's faunal zonations until the late 1970s. By that time terrestrial magnetostratigraphy, in its infancy, was being applied to Siwalik rocks. Siwalik Proboscidea were studied by Osborn (1929) and there after Sarwar (1974) worked out systematic investigation of Siwalik Proboscidea. 26
Faunal assemblages were explored (Pilbeam et al., 1977; Moonen et al., 1978) and phylogenetic investigation of each taxa were worked out and published (Dhem et al., 1958a, b, 1963; Hussain, 1971; Heissig, 1972; Jacobs, 1978; Tassey, 1983; Pickford, 1988) and subsequently, geochronologic data of Siwalik strata were compiled. The results of stratigraphy of Lower Siwalik and Middle Siwaliks were shown by Pilbeam et al. (1977) estimating chronologic range of each stage as Kamlial (before 13 Ma), Chinji (11-13 Ma), Nagri (9-10 Ma), Dhokpathan (6.5-9 Ma). The chronostratigraphy of Upper Siwalk formations was inferred by Opdyke et al. (1979) such as Tatrot (before 2.47 Ma), Pinjor (after 2.47 Ma). Lower and Middle Siwaliks were chronicled in Miocene (Pilbeam et al., 1977) while Upper Siwaliks was placed in Pliocene (Opdyke et al., 1979). In Potwar Siwaliks, one may laterally trace bands of outcrops for tens of kilometers (Pilbeam et al., 1979; Barry et al., 1980; Behrensmeyer and Tauxe 1982) for reconstructing biostratigraphic sequences (Barry et al., 1982; Raza, 1983). Stratigraphic level of fossiliferous localities was measured relating to marker horizons. Barry et al. (1982) had alleviated the nomenclatural problems of Siwalik rocks and fossils proposing a succession of four biozones for the Lower and Middle Siwalik sub-groups to replace Pilgrim's faunal zones. They reconstructed a biostratigraphic framework on the basis of faunal interval zones and calibrated with paleomagnetic polarity time scale (Johnson et al., 1982; Opdyke et al., 1979; Tauxe and Opdyke 1982). Faunal Interval zones were referred to correlate isolated fossiliferous deposits to the biostratigraphy of Siwaliks (e.g., West et al., 2010). Johnson et al. (1982, 1985) studied that the base of Potwar Siwalik sequence estimated to be >18.3 Ma, and on the top to be < 0.6 Ma. In this ca. 18 Ma interval, they identified thirteen orders of mammalian fossils. Despite of the ordinal diversity, most of the Siwalik mammals were found belonging to three orders: rodents, artiodactyls, and perissodactyls only. Flynn (1986) explored the anagenetic evolution of the rodent Kanisamys studying the succession among three species. De Bruijn et al. (1989) hypothesized the anagenetic evolution for two of the rodent species. They exemplified that the candidates for anagenetic evolution other than rodents were the Giraffokeryx and Dorcatherium. Systematic investigation of geological context of fossil localities of Potwar Siwaliks commenced with the research work of Behrensmeyer and Tauxe (1982), 27
Raza (1983), Badgley (1986a, 1986b), and Behrensmeyer (1987). Raza (1983) studied a sample of 12 localities in the Chinji formation considering associations of dominant lithology and lithofacies and relating them to the taphonomic profiles of fossil collections from the same localities. Behrensmeyer (1987; 1988) studied the fossiliferous localities within the fluvial architecture of Siwaliks at larger scale (resolution levels 3 and 4) and developed collaboration with Tauxe and Badgley (1988) for studying of isochronous deposits and fossiliferous localities at “U” level from the Dhok Pathan Formation. Flynn et al. (1990a) interpreted the Siwalik record as a representative of the geo- chronologic profile of the South Asian biogeographic province, and then compared with records from the other geographic regions to show the degree of faunal change at global scale. They predicted that the Ngorora (a part of Baringo) fauna of East African showed stability over its range, as 12 to 11 Ma interval was a time of faunal stability in the Potwar Siwaliks. Jacobs et al. (1990) further analyzed rodent fauna and extended and added field data from 1988 and 1989 to estimate species longevities. They determined an average of species longevity of 1.8 Ma and thereafter 2.2 Ma for the rodents sampled from 16 to 7 Ma interval. They compared the species longevities and revealed that earlier part of Siwaliks (prior to 9.5 Ma) documented long lived taxa with less intense turnover whereas very intense faunal turnover and short species ranges during Late Miocene was driven by paleoclimatic change. They also noted the large differences in mammalian species richness and their relative abundance between bio-stratigraphic levels. Morgan’s work, in the 1990s, on the paleoecology of mammalian fossil faunas of Chinji formation incorporated detailed taphonomic investigation of 27 fossil localities and classified them geologically (Morgan, 1994; Badgley et al., 1995). Morgan’s work was based on collaborative work with Behrensmeyer, S. M. Raza, C. Badgley, J. Kingston, and J. C. Barry highlighted significance of the floodplain or “secondary” channel from the Chinji and Dhok Pathan Formations. Badgley et al. (1998) analyzed the microvertebrate localities of Siwaliks from one stratigraphic interval from the Chinji formation, inferring that these were the result of biological agents that accumulated the bones in floodplains. Most of the fossil localities of Potwar Siwaliks document similar kinds of mammalian fossil assemblages (Barry et al., 1982; 1990), whether temporally successive 28
or geographically separated. This coherency in faunal association may reflect paleocommunity stability over substantial time intervals (Flynn et al., 1990a). The magnetostratigraphy of Potwar Siwaliks correlating temporal control on fossil localities with multiple paleomagnetic sections was worked out by Flynn et al. (1990a) and thereafter correlated these localities to those of east African biostratigraphic section and consequently Potwar Plateau displayed the classical example of sequence of precisely or well dated faunal zones in a single bio-geographic province in terrestrial settings. Having constructed the chronostratigraphic framework, different workers focused on investigation of faunal turnover patterns as baseline for understanding of paleoecological and evolutionary processes. Barry et al. (1991) worked out that the mammalian faunal turnovers in Pakistan Siwaliks occurred due to immigration, local extinction, cladogenetic speciation and anagenetic evolution. Pickford (1988) investigated change in mammalian fossil fauna of Siwaliks by comparing faunal associations on the basis of presence/absence of taxa. It may also be studied using biostratigraphic ranges of taxa to determine numbers of appearances and disappearances through particular intervals and rate of change as well (Barry et al., 1991). Barry et al. (1990) investigated a portion of fossil record of Siwaliks focusing on artiodactyls and rodents remains as representatives of ancient mammalian faunas spanning from 16 to 7 Ma and studied faunal composition on the resolution of 0.5 Ma intervals. Consequently, they calculated fluctuations in faunal diversity based on species richness and species turnover patterns at different time intervals during Middle to Late Miocene. This lead to the findings that the Late Miocene species turnover contributed mainly by the rodents, however both artiodactyls (with major emphasis on bovids) and rodents remains contributed to 9.5 Ma extinction event as well as to the earlier middle Miocene (13.5 Ma) maxima. Barry et al. (1991) linked faunal interchange with mammalian fossil diversity in Siwaliks. They also studied trends concerning increase in body size among ruminants. Barry et al. (1995) determined relative biostratigraphic positions of fossiliferous localities and interpolated the localities into magnetostratigraphic framework constructed by Opdyke (1979), Johnson et al. (1982), Tauxe and Opdyke (1982), Johnson et al. (1985) and Kappelman (1986) merely based on faunal superposition. 29
Barry et al. (1990, 1995) had utilized interval quality scores and completeness index for controlling biases that could affect faunal turnover inferences such as changes in depositional environment and fossil collection and subsequently have incorporated Koch’s (1987) method to estimate range extensions of fossil species from their occurrence data.ez After correction for biases, Barry (1995) statistically found significant maxima of first and last appearances during the intervals ranging from 13.5-12.5 Ma, and 8.5-8.0 Ma. Jacobs and Downs (1994) and Flynn (1982) studied gradual morphological change among muroid lineages. Morgan et al. (1994) investigated changes in body size of Siwalik large herbivorous mammals and rodents recorded from the Miocene, and showed that size changes started with the first appearance of hipparionine horses at
ca.10.1 Ma and the first establishing of C4 grasses at ca. 9.4 Ma. Taxonomy and distribution of bovids of Siwaliks was investigated by Akhtar (1992) and that of family suidae by Ahmad (1995). Morgan et al. (1995) further concluded that the gradual increase in their size may be attuned to more open habitats/conditions which became more accelerated between 9.0 and 8.5 Ma intervals. They explored trends in body size in different groups of small and large mammalian taxa. Barry and Flynn, (1990) and Flynn et al. (1995) considered muroid rodents since their first and last appearance data are well documented in Siwaliks. They show fairly rapid rate of speciation and even a slight hange in climatic appears to influence their abundance (Boyce, 1979, 1988; Van Dam, 1997; Van Dam and Weltje, 1999). Jacobs and Downs, (1994) and Flynn et al. (1995) have found that the muroids are supposed to have originated and progressed through anagenetic and cladogenetic patterns of evolution in the Siwaliks. Barry et al. (1995) interpreted that the Late Miocene decline in mammalian faunal diversity and increased species turnover in faunas of Siwaliks could be coincided with global conditions of increased aridity and paleo-seasonality. Gunnel et al. (1995) examined change in paleocommunity structure and composition in Siwaliks. Flynn et al. (1998) superposed faunas of Siwaliks provide a test case for examining the succession of paleocommunities spanning between 18 to 6 Ma. Barry et al. (2002) presented synergistic interplay of faunal and paleoenvironmental change during the Late Miocene (Middle Siwaliks) of northern Pakistan based on ecomorphic, lithologic and isotopic evidences. Nelson (2003) explored 30
the paleoenvironment of two different intervals of Late Miocene in association with faunal change as a major cause of extinction of Sivapithecus by incorporating microwear and stable isotopic analyses of its contemporaneous taxa of suids, small bovids, giraffids, tragulids, and anthracotheres. Patnaik (2003) explored the paleoecology and paleoclimatology of Upper Siwaliks (5.5-0.7 Ma) based on microfossil assemblages from India and he tied the described Pliocene sites to well-studied, palaeomagnetically and fission track radio-metrically dated sections. He integrated invertebrates, microvertebrates, and palynological data of Upper Siwaliks with that of Miocene deposits of both India and Pakistan to understand the shift in paleoclimate and paleoecology across the Mio-Pliocene epochs. Finally, he discussed the relative abundance of muroid rodents who replaced the cricetids in the Late Miocene and their diversification in the Late Pliocene and explored palaeoclimatological inferences. Cheema (2003) studied the phylogeny and evolution of murine rodents from Potwar Plateau and Azad Kashmir mainly focusing on zoogeographic diversification and biostratigraphic implications. Taxonomy of Swalik cervids, horses and carnivores was worked out by Ghaffar (2005) and published research papers on each of the studied group. Bhatti (2005) studied taxonomy and biogeography of Siwalik giraffids based on morphometric analysis. Systematic investigation on Siwalik Tragulids was worked out by Farooq (2006). Khan (2006) done systematic work on ruminants of Hasnot area, Late Miocene of Pakistan and published his work on boselaphines and Dorcatherium taxa of Hasnot area. Belmaker et al. (2007) worked out the mesowear-I analysis of ungulates from the middle to Late Miocene of the Siwaliks of Pakistan and to explore their dietary habits and paleoenvironment. Badgley et al. (2008) studied the impact of regional and prolonged paleoclimatic forcing on trophic structure and species richness of Miocene mammalian communities. Morgan et al. (2009) investigated Miocene vegetation gradient in the Siwaliks of Pakistan on the basis of lateral trends in stable carbon isotopic ratios. Khan et al. (2008) published their work on reduncine fossils from Upper Siwaliks of Tatrot. Systematics of boselaphines, chalicotheres (Middle Siwaliks), Proamphibos (Upper Siwaliks) and mammalian remains (Chinji Formation) of Siwaliks of Pakistan was also studied by Khan et al. (2009 a, b, c, d). Taxonomy of Siwalik Rhinoceroses was studied by Khan (2009) and published his work in peer reviewed local and international journals 31
on different taxa of rhinos and ruminants. Mammalian fossil fauna of Chinji Formation of Dhok Bin Mir Khatoon was studied by Samiullah (2010). The systematic study of artiodactyles of DhokPathan Type locality from Middle Siwalik and of Bison remains from Upper Siwaliks of Pakistan was published by Khan et al. (2010a, b). Khan et al. (2011a, b) and Khan et al. (2011) published research articles on new collection of hipparionines from Middle Siwaliks, ungulate remains from newly discovered fossil locality of Lava, Chinji formation, and Sivatherium from Upper Siwaliks of Pakistan. Khan and Akhtar (2011) studied taxonomy of Proamphibos kashmiricus from the Pinjor Formation of Pakistan. True Ungulates from Nagri Type locality, and Mammalian remains from Chinji Formation were also investigated by Khan et al. (2012a, b). Systematic investigation of Hydaspitherium cf. megacephalum and Giraffa punjabiensis from Middle Siwaliks of Pakistan was worked out by Bhatti et al., (2012a, b). Generalized interpretation of paleoecology of each studied taxon was also discussed in their endeavors. Tragulids from Chinji Formation and antelopes from upper Siwaliks (Soan Formation) was studied by Khan et al. (2013, 2014). Singh et al. (2013) worked
out the drivers that triggered the cascade of changes from forest to transitional C3/C4 vegetation followed by origination and expansion of C4 grasslands in the Siwaliks of Pakistan, India and Nepal. A systematic, biostratigraphic, and paleobiogeographic reevaluation of the Siwalik hipparionine Horse assemblage from the Potwar Plateau, Northern Pakistan (10.4-6 Ma) was studied by Wolf et al. (2013). They assessed the dietary preferences of hipparionines on the basis of mesowear analysis. They found that the progressive shifts in mesowear signatures indicated decreasing proportions of high and sharp cusps and increasing of low and blunt cusps during Late Miocene. They also observed a trend toward a decreased browse component and an increased graze component in the diet of Siwalik hipparionines while retaining their mixed feeding strategies. This trend of increasing low and blunt cusps suggest the inference that forests and woodlands were replaced by open grasslands with warm season grasses during the Late Miocene.
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CHAPTER 3 MATERIALS AND METHODS
3.1. MESOWEAR TYPE I AND II Mesowear method was applied on molar teeth of a giraffid; Giraffokeryx punjabiensis, two boselaphines; Selenoportax sp., Tragoportax sp. and an antelope; Gazella lydekkeri collected from different fossil localities of middle to Late Miocene of Pakistan (14.2-5 Ma). Specimens (molar teeth) of Giraffokerx punjabiensis from the middle Miocene of Pakistan Siwaliks were studied. Late Miocene Boselaphine taxa were also examined for mesowear analysis considering cusp shape and cusp relief (Figure 3.1) following the method outlined in Fortelius and Solounias (2000). The mesowear method determines average life long dietary adaptations on the basis of two variables namely cusp shape and occlusal relief. Both variables were determined by direct observation and the percentage of teeth with high/low cusps and sharp/round/blunt cusps was calculated for each species. These variables were then plotted against HI, as recommended by Fortelius and Solounias (2000) (Figures 3.1, 3.2). The cusp sharpness and degree of relief are dependent variables. Higher occlusal relief tends to be sharper as compared to low relief cusps and cusps with zero relief are obviously blunt. Hence, old mesowear (Mesowear Type I) has been treating as a single variable (Mesowear Type II) during which cusp apices are assigned to stages along a continuum ranging from the sharpest cusps with the highest relief to the bluntest cusps with the lowest relief. Mesowear scorings were standardized by correlating them with a mesowear “ruler” following Mihlbachler et al. (2011) (Figure 3.3). Having recorded the mesowear scorings, the percentages for each variable were calculated which in turn were examined through hierarchical cluster analyses using PAST vs 14 software to assess the dietary classification of each taxon. The purpose of incorporation of this mesowear study of Miocene ungulates is primarily to evaluate the microwear and other ecomorphic results as baseline for strengthening the evidences favoring the changes in paleoclimate and paleovegetation (forest to savannas and grasslands) during Miocene Siwaliks of Pakistan.
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3.2. MESOWEAR TYPE-III (ENAMEL BAND-2 MESOWEAR) 3.2.1. DESCRIPTION OF THE NEW METHOD OF MESOWEAR: MESOWEAR TYPE III In this study, the mesowear of band 2 was investigated in detail hoping to achieve better resolution than that from band 1 (mesowear I and II). Browse and graze experiments using adult goats (Capra hircus) were devised. Before the experiment, the goat population most likely fed on on leaves of Malia azedarach, and Morus alba (mulbery tree) and grasses such as Cynodon dactylon (khabal grass), Sorghum helepense (johnson grass) and Echinochloa colona. The experiments were performed in two phases. During first phase, five goats were placed in an enclosure and were fed browsing plants; Malia azedarach, Morus alba and Zizyphus jujuba for duration of 18-20 days. Five different goats were placed under the same conditions and were fed grazing plants; Cynodon dactylon, Sorghum helepense, Echinochloa colona for the period of 18-20 days. In the second phase, eight goats were utilized. One goat was subjected to browsing for 10 days. After that period, the goat was euthanized and the skull and jaw were skeletonized. The next goat was subjected to 20 days of browsing, the next one for 30 days, and the last one for 40 days, and the same procedure was performed. This was repeated for four additional goats that were subjected to grazing on Cynodon dactylon, Sorghum helepense, Echinochloa colona, Dichanthium annulatum, Digitaria violascens and Echinocloa colonum, keeping all other conditions the same. Fecal materials and rumen contents of all the animals were preserved in formaline solution. In order to do this properly, band 2 from an occlusal view of extant wild specimens of browsers, grazers and mixed feeders was evaluated. Dental mesowear-III pattern of goats was examined and compared with the wild ruminants. Finally the results were applied to a selection of 17 fossil species of ungulates from Pakistan and Late Miocene giraffids from China. A new scoring scale of only 4 wear states (Figure 3.6) has been developed. Scoring was done on the occlusal anterior part of band 2 separately from the posterior and subsequently averaged the two. The mesowear of all specimens were evaluated qualitatively by observation with a stereo-microscope. From observation of the teeth of known wild extant grazers and browsers, a method was developed using enamel band 2 to interpret the diet of the individual. In the following paragraphs, a morphocline 34
sequence going from the ideal browser to the ideal grazer is described. This morphocline has been divided into 4 scores, although it is known that the actual wear of the teeth is a continuum. It is believed that one can use these 4 scores to extrapolate the diet of an individual. For band 2, there are two sides (the mesial side and the distal side), separated by a midpoint (termed j for junction). The mesial side has different mesowear from the distal side using mesowear III (to be described). The contraction of the temporalis muscles during occlusion and cutting of vegetation forms the mesial side and the distal side on both the paracone and the metacone. The anterior side is the leading edge, considering the direction of the bite force. At the mesial side, there is higher force because the mandible is pulled back against the upper mesial side, which is tilted. The leading side is often a flatter edge and is subjected to more attrition. The distal side trails during the cut and most likely has a weaker attritional contact. Due to this difference in forces during mastication, the mesial side and the distal side often have different mesowear III and are therefore scored separately. The two scores are averaged to form a collective evaluation for the entire band. 3.2.2. DEVELOPMENT OF SCORING SCALES Examination of teeth of goats and other species suggests that two new scoring scales concerning browsing-grazing mesowear sequence can be developed. 3.2.2.1. BAND-2 SCORING SCALE Score 1; is the ideal browser morphology for the mesial or the distal side of the cusp.The facet is one large, planar surface. The edge of the surface is relatively sharp, with no gouges. Score 2; The surface is no longer one large, continuous unit, but is broken up into two to four smaller sub-facets. The edge not as sharp as in score 1. The edge has gouges, primarily on the buccal side. Score 3; The surface is similar to score 3 but is broken up into more sub-facets and has more gouges. There is also rounding between the sub-facets and on the edges. Score 4; is the ideal grazer morphology for the mesial or the distal side of the cusp. The edges of the enamel band are rounded instead of sharp with no gouges or sub-facets, and the surface forms a uniform arch. The scoring is shown in figures 3.4-3.6. 35
3.2.2.2. J-POINT SCORING SCALE The two sides of the cusp have a mid-point where they merge. This mid-point is named as the junction point or J point. The wear of the j point has been scored separately from the wear of the sides. Score 1; When j is very sharp and well defined, I score it as 1. Score 2; A j with score 2 still has the sharp edge but contains one or two gouges and sometimes contains a small facet. Score 3; A j with score 3 is more rounded but is still visible. Scores 2 and 3 are intermediate stages. Score 4; When j is completely absent and the 2 sides form one continuous surface, I score it as 4 (Figures 3.4-3.6). The band 2 of the metacone of the upper right second molar was used when scoring all goat teeth. For other species, band 2 of the metacone was used when it was available, but when that region was damaged; band 2 of the paracone was considered for scorings. For the extant ruminants, the specimens were used from the American Museum of Natural History. The teeth were molded and cast. The browsing animals used were: Okapia johnstoni (N=11), Giraffa camelopardalis (N=15), and Alces alces (N=13). The grazing animals used were: Ourebia ourebi (N=7), Kobus ellipsiprymnus (N=10), and Chonnochaetes taurinus (N=12). The mixed feeders used were: Cervus canadensis (N=4) and Grazella granti (N=17). For the extant ruminants and remains of Giraffokeryx punjabiensis, specimens from the American Museum of Natural History were used. The teeth were molded and cast. For the fossils, specimens of remains of boselaphines and hipparionine horses housed in Paleontology Laboratory, Department of Zoology, Government College University, Lahore, were used. Specimen’s of Late Miocene giraffids from China were also incorporated in these studies which are housed in the following museums: Paleontological Institute of Uppsala (PIU), Institute of Vertebrate Paleontology and Paleoanthropology at Beijing (IVPP), and the Hezheng Paleozoological Museum. The mesowear of all specimens were evaluated qualitatively by observation with a microscope. The t tests were run between browsers and grazers and then a t for each unknown against browser and against grazer. Now if the p-value is strong, then the unknown in question will be either a grazer or a browser. If it is weak it will be a mixed feeder. 36
3.3. MICROWEAR METHOD A sample of 128 molar teeth attributed to twelve (12) fossil taxa of mammals was selected for exploration of paleoecology of Late Miocene of Siwaliks applying dental microwear method. The selected taxa represent most of the major groups of mammals that include two suids; Propotamochoerus hysudricus and Hippopotamodon sivalense, five bovids; Elachistoceras khauristanensis, Pachyportax latidens, Selenoportax vexillarius, Tragoportax sp., Miotragocerus gluten, a tragulid; Dorcatherium majus, a giraffid; Giraffokeryx sp., an anthracothere; Merycopotamus nanus, a hipparionine horse; Hipparion sp. and two rhinos; Brachypotherium fatejhangense, Brachypotherium perimense. The purpose of this microwear study of ungulates is to evaluate the changes in dietary composition of these taxa that strengthen the hypotheses concerning the evolution of paleovegetation from forest to savannas and grasslands and their associated paleoclimate and paleoenvironment during Middle to Late Miocene Siwaliks of Pakistan. This study followed the detailed procedure for molding; casting and assessing specimens mentioned in Solounias and Semprebon (2002) and Semprebon et al., (2004). Casts of epoxy steel of fossil teeth were analyzed under light stereo-zoom microscope (Olympus SZH 10) at 35X magnification by Nikos Solounias (New York Institute of Technology College of Orthopathic Medicine) and was termed as NS magnification. Counting of microwear characters was done in a 0.5 x 0.5 mm area using occular micrometer/reticle. Casts of same fossil specimens were also analyzed under stereo-zoom microscope (SZM 405-S2, HT Company, UK) by present author at 35X and was termed as MT (Muhammad Tariq, s) magnification. Microwear results of NS magnification and MT magnification were compared for drawing dietary inferences (Appendix 4). Microwear data on Hippopotamodon sivalense, Propotamochoerus hysudricus and Dorcatherium majus from Siwaliks of Pakistan compiled by Nelson (2003) were also incorporated for comparative reasons with the purpose to work out more reliable dietary evaluations of the studied taxa. The measured microwear features are comprised of number of scratches and number of pits. Pits are those microwear features that are circular or semicircular in shape and show length: width ratio below 4:1. Large pits are wider and deeper but appear dark and less refractive whereas small pits are comparatively shallow, appear bright and shiny hence refract light easily. Scratches reflect length: width 37
greater than that of pits. They are elongated scars with parallel, straight sides and may be sub-categorized as fine or coarse. Fine scratches are narrow, and barely etched into the enamel surfaces. Coarse scratches are wider and more obviously etched into the enamel surface. In ungulates, most dental scratches indicate the direction of jaw motion hence these scars are not erosional. Cross scratches are oriented somewhat antero-posteriorly in the mouth and are probably produced as stems are stripped off by the animal. Gouges are large pits but are very irregular in shape. Cross scratches are those characters that run perpendicular to each other (Solounias and Semprebon, 2002) (Figure 3.10 A, B, C, D, E, F). Microwear data for each species are listed in appendix 2. Microwear pattern of selected fossil species were compared to microwear signatures of their modern analogues with known habitats and diets to evaluate their ecological role. For ease of comparison, microwear database of living ungulates compiled by Solounias and Semprebon (2002) and by Nelson (2005) have been incorporated in this study. Bivariate plots dully marked by dietary morphospaces of living ungulates were designed in PAST (paleontological statistics) 2012 that reveal distinct morphospaces indicative of browsing, grazing, and mixed diets. The taxa were also assigned to their respective trophic groups based on ranges of scratch distribution (Table 4.7). The radical dietary classification of ungulates was worked out on the basis of Microwear Index (MI). The microwear index (MI) was calculated as the average number of scratches divided by average number of pits. As a standard, an MI > 1.5 is indicative of a grazing diet and an MI < 1.5 represents a browsing diet (MacFadden et al., 1999b). Additional conclusions were drawn by comparing the analyzed tooth to a database of the microwear pattern of extant analogues with observed known diets. Mann-Whitney and student t-test were applied to test for significant differences in microwear features. Statistical analyses were run on SPSS version 15.0 with significance set at p< 0.05. 3.3.1. ESTABLISHING THE MORPHOSPACES The bivariate plots of the average number of scratches versus average number of pits for extinct and living herbivores have developed distinct morphospaces indicating browsers, grazers and mixed feeders (Figures 4.23-4.29). The boundaries of dietary morphospaces of extant taxa are demarcated by shaded areas in these plots. Three dietary groupings of livings are represented: (1) leaf browsers (LB)__ relatively low number of 38
scratches but variable numbers of average pits; (2) grazers (GR)__relatively high number of scratches while low number of pits; (3) fruit browsers (FB)__ generally show moderate numbers of pits and scratches as compared to other two extreme dietary groups. 3.3.2. QUANTITATIVE CATEGORIZATION OF AVERAGE MICROWEAR COUNTS EXAMINED IN INDIVIDUAL SPECIMENS Typical leaf-browsers have high/more disparate average number of pits and low average number of scratches with most of being fine whereas fruit-dominated browsers bear intermediate number of pits and scratches. Typical grazers reflect high average number of scratches and low average number of pits. Mixed feeders normally show values of scratches and pits overlapping to those of grazers or browsers depending upon the relative amount of grass or browse consumed regionally or seasonally.Seasonal mixed feeders indicate texture of scratches and average number of scratches among individual of species by displaying bi-modal distribution when average number of pits versus average number of scratches plotted (Solounias and Semprebon, 2002). One may distinguish mixed feeders from grazers and browsers by simply calculating the % age of individuals of the studied taxon that have number of scratches in low range (0-17 % scratches). Browsers show high percentages of specimens with low number of scratches ranging from 72-100%, on the other hand, grazers represent relatively few specimens placing in low range of scratches (0-22%) while mixed feeders show the range of 20-70% with low range of scratches (Semprebon and Rivals, 2007; Solounias et al., 2010). 3.4. HYPSODONTY METHOD The general profile of dental morphology of mammalian fossil fauna and ecomorphic information of 160 taxa of ungulates from Siwaliks of Pakistan were taken as sole evidence for paleoenvironmental reconstruction. A sample of 365 specimens of ungulates was selected for hypsodonty analysis. The selected specimens were found belonging to three orders, eight (8) families and 67 taxa. Ecomorphic data of 34 taxa were collected from tha samples housed at Paleontology Laboratory, Department of Zoology, GC University, Lahore whereas ecomorphic data of 33 taxa of ungulates were compiled from published sources. Among 67 studied taxa, 55 taxa were treated at specific level and 12 taxa at generic level for hypsodonty analysis following Fortelius et al. (2002, 2006), Solounias et al. (2010) and Damuth and Janis (2011). The 39
morphometrics of studied taxa were translated into ecometrics (diet and habitat etc). The metrics of unworn third lower molars were recorded with digital vernier callipers. The ranges of hypsodonty values in ungulates have been used to categorize dietary classes such as brachydont, mesodonts, and hypsodonts to infer changes in diet and habitat. Ecomorphic data of 93 taxa of Siwalik ungulates was compiled by comparing with their extant analogues. Among 93 taxa, 71 taxa were studied at specific level, 18 taxa at generic level, 2 taxa at tribe and 2 taxa at sub-familiy/family level following Barry et al. (2002); Fortelius et al. (2002, 2006); Badgley et al. (2008). The inference of paleohabitats or regional paleoenvironment was corroborated with the hypsodonty values and habitats of extant taxa. Measurements of crown height and length were taken on teeth that were relatively unworn, and represent the greatest height and length for each tooth. The specimens that reflect similarities in taxonomy and in morphology and ecology between the extant species have been treated at generic level. The ecomorphic information were also downloaded from paleobiology database (2013) and were compared with existing taxonomic data collected by various workers such as Sarwar (1974), Akhtar (1992), Barry et al. (2002), Gaffar (2005), Bhatti (2005), Farooq (2005) Khan (2006), Khan (2009) on Siwalik ungulates. The body weights of extinct taxa of mammals were explored and evaluated by comparing with the body weights of their extant analogues tabulated in Janis (1988). 3.4.1. DIETARY AND HABITAT CATEGORIZATION OF SIWALIK MAMMALS The dietary and habitat categories of mammalian fossils have been incorporated in this study following Hofmann and Stewart (1972), Janis (1988), Damuth and Janis (2011). A. Omnivores (O); The organisms taking in of considerable amount of herbage (predominantly rootles along with fruits and other types of vegetation) and animal materials in their diet are recognized as omnivores. This category is used only in certain taxa of suines/suoides. B. Browsers (B); The organism taking less than 10% of grass and >90% dicotyledonous herbage in their diet are called browsers. 40
(i) Regular Browsers (BB); The browsers taking less than 10% of grass and >90% dicotyledonous herbage in their diet which is equivalent to “tree and shrub foliage eaters” of Hofmann and Stewart (1972). (ii) High level browsers (HB); The browsers taking <10% of grass and >90% dicotyledonous herbage in their diet and use to be feeding above the ground level invariably are categorized as “high level browsers” “HB” e.g. giraffes, gerenuk, moose (Damuth and Janis 2011). (iii) Selective Browsers (SB); The browsers that habitually incorporate the considerable amount of fruit, buds and berries in their diet (and preferablly using young to mature leaves) are classified as “selective browsers” “SB” (Damuth and Janis 2011) = “fruit and dicot foliage selectors” (Hofmann and Stewart 1972). C. Mixed feeders (MF); The organisms taking >10% and <90% grass in their diet are treated as mixed feeders. (i) Mixed feeders in closed habitat (MFC); Mixed feeders in closed habitats (MFC) are inhabited in forest or woodland and bushland habitats. This category is broadly (but not entirely) equivalent to “intermediate feeders who prefer shrubs, forbs and tree foliage” of Hofmann and Stewart (1972). (ii) Mixed feeders in open habitat (MFO); The organisms taking >10% and <90% grass in their diet are considered as mixed feeders in open habitats. They are found in savannas (treed grasslands) or prairies (treeless grasslands). This category is broadly (but not entirely) equivalent to “intermediate feeders who prefer grasses” of Hofmann and Stewart (1972). D. Grazers; The organisms taking > 90 % of grass in their diet are known as grazers. (i) Fresh grass grazers (FG); The grazers grazing near water or in edaphic habitats are classified as fresh grass grazers (FG). (ii) Dry grass grazers (GG); The grazers grazing in dry regions [lump together “dry region grazers” and “roughage grazers” of Hofmann and Stewart (1972)] are categorized as dry grass grazers (GG).
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3.4.2. MEAN HYPSODONTY Mean Hypsodonty Patterns explore the conditions of the vegetation which may be nominated as “generalized water stress” depicting the humidity-aridity gradients. Mean Hypsodonty has direct proportion with arid conditions and inverse proportion with the humid environment. This implies that the higher value of mean hypsodonty is indicative of more arid conditions, whereas low hypsodonty value represents more humid environment (Fortelius et al., 2002, 2006; Eronen 2006). The investigation was constrained to localities of Pakistan Siwaliks, and only large mammals were selected and classified as plant-eaters or plant-dominated omnivores (suids) for analysis. Localities with at least 2-3 taxonomically resolved species of true ungulates were included in the analysis. Three classes of hypsodonty were assigned scores of 1) brachydont, 2) mesodont and 3) hypsodont. The mean hypsodonty value in every locality was calculated by averaging its herbivore scores. The possible range of mean hypsodonty is thereby constrained to values between 1 and 3. Ranges of mean annual precipitation estimations and mean annual temperatures were translated from mean ordinated hypsodonty values. The mean hypsodonty and precipitation values were plotted on maps using interpolated contours following Fortelius et al. (2002, 2003and 2006) during which modern maps were used as a background for these patterns. GIS maps portraying pictorial display of ranges of mean ordinated hypsodonty and annual precipitation from Siwalik region were prepared in Arc- Map Info 10.
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FIGURES
Figure 3.1; Tooth Mesowear displaying selection of three types of apices (cusps)
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Figure 3.2, Example of Mesowear-I based on two variable named cusp relief (high, low) and cusp shape (sharp, round and blunt). The dotted lines represent the height of occlusal relief (source; Kaiser and Solounias, 2003).
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Figure 3.3 A; Mesowear “ruler” designed for dental mesowear scorings of equid remains, comprising of seven (0-6) cusp apices of Equus cast onto an epoxy surface. The mesowear “ruler” is utilized by matching the dental cusp apex of a specimen of fossil species with one of the seven (0-6) reference cusps to assign and standerdize it a score of 0–6 (source; Mihlbachler et al., 2011).
Figure 3.3 B; The mesowear “ruler” applied to a fossil bovid-Urmiatherium. 45
Figure 3.4; Pattern of Mesowear-Type III of a typical browsing goat. 46
Figure 3.5; Pattern of Mesowear Type III of a typical grazing goat.
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Figure 3.6; The figure is a model made in clay portraying coding system of Mesowear Type III depicting a new development of browsing-grazing mesowear sequence. It summarizes the general findings. 48
Figure 3.7A; Mesowear-2 of browsing goats; B10 stands for browsed for 10 days, B20 browsed for 20 days. i=posterior band, ii=anterior band.
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Figure 3.7B; Mesowear-2 of browsing goats. B30 stands for browsed for 30 days, B40 browsed for 40 days. i=posterior band, ii=anterior band.
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Figure 3.8A; Mesowear-2 of grazing goats; G10 stands for goat grazed for 10 days, G20 grazed for 20 days. i=posterior band, ii=anterior band
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Figure 3.8B; Mesowear-2 of grazing goats; G30 stands for goat grazed for 30 days, G40 grazed for 40 days. i=posterior band, ii=anterior band.
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Figure 3.9 A-E, A= Mammalian molar demarcated with second band of paracone where all microwear molds and cast preparation is focused. B= Preparation of mold with applicator gun. C= stereomicroscope (Olympus SZH 10). D= Double head fiber-optic light source and Plexiglas stage used for reflected illumination from table surface. E=Typical mold surrounded by wall of lab putty to hold fluid epoxy and epoxy cast of the crown surface (Source; Solounias and Semprebon, 2000). 53
Figure 3.10, Generalized SEM photomicrographs at 500X. A= Surface showing small and large pits. B= Large puncture like seed pits, C= Mostly finelyscratched surface. D= Surface showing a gouge and cross scratches. E= Coarse scratch and fine scratches. F= Mixture of scratches (source; Solounias and Semprebon, 2000).
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CHAPTER 4 RESULTS 4.1. MESOWEAR TYPE I, II
4.1.1. Giraffokeryx punjabiensis: The mesowear signatures of Giraffokeryx punjabiensis reflect affinities to both browsing and mixed feeding extant analogues. The living analogues for browsing are a giraffid; Giraffa camelopardalis, and a cervid; Capreolus capreolus. The extant analogues for mixed feeders are Tragelaphus imberbis and Procavia capensis (Appendix 10). Mesowear data for each molar are given in appendix 7 and absolute and relative mesowear scorings of studied specimens are given in Table 4.1. The Hypsodonty Index (HI) for Giraffokeryx punjabiensis calculated here is 1.31 ± 0.06 (n=7). The purpose of incorporation of hypsodonty index (HI) is primarily to evaluate the mesowear results and to compare to its extant analogues i.e. Giraffa camelopardalis, Okapia johnstoni for drawing ecomorphic inferences. The HI of 1.31 ± 0.06 is utilized for G. punjabiensis in all bivariate representations. Catalogue numbers along with tooth occlusal height and width metrics are mentioned in Appendix 8. Heiratchical cluster analysis of G.punjabiensis amongst 30 list of extant browsers, mixed feeders and grazers from Foetelius and Solounias (2000) reveals that the mesowear pattern of studied species is consistent with browsing taxa i.e. G. camelopardalis, and a cervid; Odocoileus hemionus (Figure 4.2; Appendix 10). The graphical representation indicates that the G. punjabiensis may be placed within the dietary spectrum of browsers and seasonal mixed feeders (Figure 4.3-4.5 A, B, C). 4.1.2. Tragoportax sp.: The mesowear pattern of Tragoportax sp. shows affinities with browsing extant analogues. The living analogues for browsing are two cervids; Odocoileus hemionus, and Capreolus capreolus (Figures 4.6-4.7; Appendix 10). Mesowear data for each molar is given in appendix 7 and absolute and relative mesowear scorings of studied specimens are given in Table 4.2. The graphical representations indicate that the Tragoportax sp. may be placed within the dietary spectrum of browsers. The cusp shape of the specimens has 28.57% round cusps, 71.42% sharp and 0% blunt cusps. The Hypsodonty Index (HI) for Tragoportax sp. calculated here is 1.21 (n=4). The purpose of incorporation of hypsodonty index (HI) is primarily to evaluate the mesowear results and to compare to its extant analogues for drawing ecomorphic inferences. 55
4.1.3. Selenoportax sp.: The mesowear analysis of Selenoportax sp. reflects affinities with browsers. The living analogues for browsing are a giraffid; Giraffa camelopardalis, a Rhinocerotid; Dicerorhnus sumatrensis (DS), and a cervid; Odocoileus hemionus (Appendices 7, 10; Table 4.3). The bivariate plots (Figure 4.11 A, B, C) and cluster analysis also places the studied taxon within the dietary spectrum of browsers (Figure 4.10). As regards cusp shape, the specimens have 33.33% round cusps, 66.66% sharp and 0% blunt cusps (Figure 4.9). 4.1.4. Gazella lydekkeri: The mesowear study of Gazella lydekkeri reveals that it shows affinities with two mixed feeding bovids; Ovibos moschatus (Om) and Aapyceros melampus (Me) (Appendices 7, 10; Table 4.4). Cluster analysis (Figure 4.13) and bivariate plots (4.14 A, B, C) also place the studied taxon in dietary spectrum of mixed feeders. As regards cusp shape, the specimens have 41.66% round cusps, 58.33% sharp and 0% blunt cusps (Figure 4.12). Considering these fore-mentioned extinct species as a ecological models, mesowear type II of seven (7) more fossil species have been worked out that are listed alongwith Mesowear Type III (Table 4.5) which include; Pachyportax nagrii, P. latidens, E. khauristanensis, Miotragocerus gluten, Merycopotamus nanus, Dorcatherium majus and Hipparion sp. 4.2. MESOWEAR TYPE-III ANALYSIS The separation between the extant browsing and grazing ruminants was statistically significant. Two sample t-tests, assuming unequal variances, were performed in Matlab (Mathworks, Inc.) using the test 2 function to compare the data set. As expected the mixed feeders overlapped with the grazers and the browsers (Figure 4.15 a, b). The results of mesowear III are better than those of mesowear II (Table 4.5); they show finer discrimination. The results for the goat experiment are preliminary as the sample is small. Before the experiments, the goats started off with the mesowear III of a browser. The red line in Figure 4.17 shows a progression of the mesowear for the four goats from 10 day browsing to 40 day browsing. The teeth became incrementally flatter during the period of 30-40 days of browsing. Note the graph is made by four different goats sacrificed at the termination of that period. In other words it is not the same individual aging. The green line in Figure 4.16 shows a progression of the mesowear for the four goats from 10 day grazing to 40 day grazing. The teeth became incrementally 56
rounder for the first 30 days of grazing, and did not change during the last 10 days (Figure 4.17). Mesowear-III pattern of G. punjabiensis (average of mesowear band 2 is 1.20 and that of J-point is 1) resembles with the extant browsing giraffids; Giraffa camelopardalis, and Okapia johnstoni (Tables 4.5-4.6). Pachyportax nagrii is inferred as a browser since its mesowear III patthern is consistent with browsers (Table 4.6; Figure 4.18). E. khauristanensis is presumed to have browsing/frugivorous adaptations. Mesowear-III of Tragoportax sp. shows that the animal was browser in woodland. Miotragocerus gluten exhibit mesowear pattern of browsers whereas Pach. latidens and Selenoportax sp. are considered to be mixed feeders in open vegetation (Figure 4.19). Gazella lydekkeri reflect mesowear III signatures consistent with browsers to mixed feeders (Table 4.6; Figure 4.24). Hipparion sp. is hypsodont grazer that resembles extant equines as cluster analysis categorizes it among the grazers (Figure 4.22). Mesowear III of Dorcatherium majus (mesowear band 2= 1.58, J-point= 1.16) resembles with browsing Capra hircus and extant species of Giraffids. It may fall within the dietary spectrum of browsers and frugivores (Table 4.6; Figure 4.20). Merycopotamus nanus is inferred as browser in humid and swampy habitats (Figure 4.23). Mesowear-III analysis indicated that Paleomeryx kaupi-sanson, Paleomeryx kaupi-artenary and Giraffokeryx punjabiensis were browsers whereas five species named Schasitherium tafeli, Palaoetragus coelophrys, Honanotherium schlosseri, Alcicephalus neumaryi, Samotherium boissieri, Samotherium sinense were mixed feeders (Table 4.18). The browsing and mixed feeding taxa of giraffids from Late Miocene of China were ecologically equivalent to the lineages of Giraffa and Hydaspitherium-Bramatherium from Siwaliks of Pakistan respectively (Table 4.6, Figure 4.18). For knowing dietary inferences of extinct giraffids, t-tests were run between browsers and grazers and then a t for each unknown agaist browser and against grazer. If p-value is strong with browser and weak with grazers, the organism is browser. If p-value is strong with grazer and weak with browser, the animal is grazer. If p-value is shown to be weaker with both browsers and grazers, the animal is categorized as mixed feeder (Table 4.18).
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4.3. MICROWEAR ANALYSIS 4.3.1. DIETARY ADAPTATIONS OF UNGULATE REMAINS BASED ON MICROWEAR When fossil species are plotted outside the grazing and browsing morphospaces of the living species (Figures 4.21-4.27), their dietary preferences may either be mixed feeders or browsers or grazers who are truly placed outside the boundaries of extant morphospace. This is very interesting as it suggests that the paleoecology of middle and late Miocene of Siwaliks of Pakistan was different from any present day ecology. 4.3.1.1. Suidae: Two fossil species of suids; Propotamochoerus hysudricus and Hippopotamodon sivalense are fruit dominated browsers as their scratch distribution range falls within the dietary morphospace of fruit browsers (FB) (Figure 4.25, Table 4.7). When H. sivalense and P. hysudricus were compared to their three extant analogues named Babyrousa babyrousa, Potamochoerus porcus and Hylochoerus meinertzhazeni. P. porcus is an African red river hog with dietary composition including fruit as major component and rootles as second one. B. babyrousa (the pig deer from Southeast Asia) includes fallen fruits in its diet only. Hylo. meinertzhazeni is a giant African forest hog and feeds on foliage leaves primarily of shrubs, sedges, short grasses and herbs, little to no fruit (Kingdon, 1997; Nowak, 1999; Appendix 11). Microwear pattern of both H. sivalense and P. hysudricus, reflect a large number of pits with large pits and dominant mixed scratches which is similar to the microwear pattern of predominantly frugivorous living pigs. Nevertheless, individual microwear assessment reveals that some specimens show very little or no fruit microwear signatures but foliovorous. The primary component of Propotamochoerus diet was fruit which may have only been abundant seasonally and during significant interim of little availability of fruit, the animal used to be foliovorous. The dietary adaptations of Propotamochoerus hysudricus reveal the marked seasonality during the late Miocene. The diet of H. sivalense was very much similar to P. porcus, followed by Hylochoerus and Babyrousa. When assessed individually, 14 specimens were frugivorous, one looked like foliovorous and eight showed mixed diet. Individual microwear assessment suggests that there are no significant differences found in number of scratches or pits over time for each species indicating no evolution in dietary behavior in these two extinct suines throughout the 58
Late Miocene (Appendix 3). Dietary preference of Propotamochoerus is suggestive of seasonal variability in availability of fruit. The Hippopotamodon was feeding on fruits but different to that of Propotamochoerus and as a consequence was less susceptible to seasonal changes. This suggests that two species were not contemporaneous. MIs and percentages of low scratch distribution range (31.9-37.5%) of both the species indicate the incorporation of browse food in their diet (Appendices 3, 4; Table 4.7). 4.3.1.2. Tragulidae: Microwear of one fossil species of tragulids was compared to the microwear of its extant analogues. The percentage of low scratch distribution range (50%) and established dietary morphospaces suggest that Dorcatherium majus is a leaf browser (Figure 4.26; Table 4.7). As is observed in modern tragulids, the microwear of each fossil species of tragulids shows high number of pits and coarse scratches that spans
all dietary adaptations except C3 grazing. The heavy wearing among the tragulids is most likely because of feeding on hard covered fruit and seeds. Dorcatherium majus, incorporated smallest browse and largest fruit components in their diets. Many duikers and small antelopes primarily feed on fresh leaves, shoots and buds but they are confined to moist forested habitats where foliage are available year round (Kingdon, 1997). Microwear Index (MI) = 1.53 by NS magnification and 1.46 by MT magnification is also indicative of incorporation of fruit as major component with browse in the diet (Appendix 4). 4.3.1.3. Boselaphini: As regards boselaphines, four fossil species have been referred for microwear studies named Pachyportax latidens, (n=5) Selenoportax vexillarius, (n=6) Miotragocerus gluten,(n=1) Tragoportax sp. (n=6) from latest Miocene (8-6 Ma). Pachyportax latidens, a large bovid, has microwear pattern (MI= 1.45 by NS magnification and MI=1.50 by MT magnification) concordant with mixed feeders in open habitat. Selenoportax vexillarius has microwear consistent with fruit and graze components in the diet (Appendix 11). MIs of studied taxa are 1 and 1.03 as revealed by NS and MT magnification respectively (Appendix 4). The dietary preference of Selenoportax sp., was predominantly fruigivorous which did not change throughout its explored biostratigraphic range (Belmaker et al., 2007). Microwear Index (NS magnification shows MI= 1.32 whereas MT magnification shows MI=1.28) of Tragoportax sp. suggests that it had been feeding on browse diet. Medium sized bovids 59
(e.g. Tragoportax sp.) were feeding on browse diet before 8 Ma and after 8 Ma incorporated fruit and graze in their diet (Belmaker et al., 2007). Miotragocerus gluten is infered as browser/mixed feeder as its MI is < 1.5 based on MI of NS magnification is 1.44 and its MI from MT magnification comes out to be 1.38 (Appendix 4; Figure 4.27). 4.3.1.4. Giraffidae: Among the giraffids, Giraffokeryx sp. is represented by two specimens (M2s) dated at about 9.5 Ma. Bivariate plot displays the studied taxon within the boundary of browsing morphospace (Figure 4. 28; Appendix 11). Its Microwear Index based on NS magnification (MI=0.55) and MT magnification (MI=0.60) also suggests that the animal has microwear pattern consistent with browsers (Appendix 4). Nevertheless, it is preliminary microwear analysis as the sample size (n=2) is very small. 4.3.1.5. Anthracotheroidae: Microwear of Mery. nanus shows more average number of scratches than average number of pits since its Microwear Index (MI) exceeds 1.5 (MI= 1.77 by NS magnification and 1.51 by MT magnification) (Appendix 4). The data reveal the grazing diet in open habitat although the sample size is very small (n=3). Bivariate categorizes the taxon as grazer because it is confined in grazing morphospace (Figure 4.29; Appendix 11). 4.3.1.6. Equidae: Hipparionine horses are among the most hypsodont taxa of late Miocene of Siwaliks. The microwear analysis (MI = 2.83 by NS magnification, MI= 2.60 by MT magnification) of Hipparion sp. (Appendix 4) from Hasnot area confirms it as grazer in open habitat. The similar results are represented in bivariate plot (Figure 4.30). 4.3.1.7. Rhinocerotidae: The microwear data on the rhinocertids are considered as preliminary since the sample size is very small. Specimen of Brachypotherium fatejhagense (n=1) is suggestive of browser whereas that of B. perimense (n=1) is considered as a grazer in dietary adaptations (Figure 4.31; Appendix 4). 4.4. HYPSODONTY ANALYSIS 4.4.1. Boselaphini: Considering Janis (1988), Damuth and Janis (2011) hypsodonty interpretations, Selenoportax sp., Pach. latidens and Ruticeros pugio may be categorized within the dietary spectrum of mixed feeders in open habitats (MFO) that are analogous to HI of Cervus canadensis and Camelus dromedaries (Appendix 12). HI=Tragoportax sp. is presumed to have been browser as Odocoileus virginianus is, before 8 Ma and mixed feeder afterwards. Miotragocerus gluten was a mixed feeder in 60
closed/semi-closed habitat (MFC) leaf and herb-eater in shrubland-light woodland habitats (Table 4.8; Figure 4.32). HI of Miotragocerus large sp. is indicative of leaf and herb-eater in shrub-land to light woodland habitats (MFO) consistent with Ozotoceros bezoarticus (MFO). The small sized (<15 Kg) boselaphine lineages; Helicoportax praecox, H. tragelaphoides, Helicoportax sp., Tragoportax salmontanus, Miotrag. salmontanus and E. khauristanensis were selective browsers/ fruigivores. Tragoceridus sp. was mixed feeder in closed habitat (MFC) before 8 Ma and its dietary habit was changed to mixed feeder in open habitat (MFO) after 8 Ma. The Sivoreas eremita, Sivaceros gradiens and Strepsiportax sp. show dietary spectra of all types of browsers to mixed feeders. Eotragus noyie is categorized as high-level browser (HB) and regular browser (BB). The rarely occurring Kubanotragus skolovi, Protragocerus gluten were browsers of all types whereas Paleohypsodontus zinensis is considered as mixed feeder in closed habitat and is analogous to Cervus nippon, Cervus unicolor equines (Table 4.8; Figure 4.32; Appendix 10). 4.4.2. Bovini: The ecometrics of Proamphibos lacrymans, Hemibos triquetricornis, Bos sp., Bison crassicornis, Bison sivalensis and Bos acutiferons indicates that the organisms were mixed feeders in open habitat (MFO) similar to that of their extant analogues whereas Babalus palaindicus (HI=2.85) and Proamphibos sp. (HI=4.53) are considered as fresh grass grazers (FG) (Table 4.10; Figure 4.33). Damalops palaeindicus was mixed feeder in open habitat and body weight (B.W=235 kg) falls within the range of modern hartebeest (125 Kg) to wildebeest (342 Kg) (Appendix 10). 4.4.3. Reduncini and Antilopini: Reduncini large species, Kobus porrecticornis, “Reduncini/ D013 species” and other reduncine taxa were fresh grass grazers as extant reduncines are fresh grass grazers (FG) and mainly associated with the wet meadows, permanent drainages or floodplains (Table 4.9). Three antelopine species Antilope subtorta, Antilope planicornis and Antilope intermedius were mixed feeders in open habitat (MFO). Gazella sp. was mixed feeder in closed habitat (MFC). The other younger member of Gazellini; Gazella lydekkeri was mixed feeder in open habitat (MFO) (Table 4.9; Figure 4.34). The cf. Prostrepsiceros vinayaki, Protragelaphus skouzesi are also categorized as mixed feeders. 61
4.4.4. Giraffids: The HI for Giraffokeryx punjabiensis (1.31) may be placed in brachydont category as explored by Janis (1988) and Fortelius and Solounias (2000). The dental morphology shows affinities with brachyodont forms. Considering Janis (1988), Damuth and Janis (2011) hypsodonty interpretations, G. punjabiensis may be categorized within the dietary spectra of all the three types of brachyodont browsers such as regular browsers (BB), selective browsers (SB) and high level browsers (HB) (Table 4.11; Figure 4.36). Furthermore, the HI of G. punjabiensis is very similar to the average for four regular browsers (BB); Odocoileus virginianus, Hylo. minertzhageni, Mazama mazama americana, Choeropsis liberiensis, two selective browsers (1.31 ± 0.06) Cephalophus dorsalis, Hyaemoschus aquaticus and three high level browsers Alces alces, Litocranius walleri, Okapia johnstoni. Having compared the HI and body size/mass of Giraffokeryx punjabiensis to the previously published HI and body size of 127 species of extant ungulates (Janis 1988), G. punjabiensis HI seems most similar to that of its multiple extant analogues i.e. one cervid, Alces alces, a giraffid Okapia johnstoni, Choeropsis liberiensis (pygmy hippos), a suid; Hylo. minertzhageni (giant forest hog). The Alces alces and O. johnstoni have been classified by Janis (1988) as high level browsers (HB), while C. liberiensis and Hylo. minertzhageni as regular browsers (BB), Cephalophus dorsalis, Hyaemoschus aquaticus as selective browsers (SB) (Appendix 12). Both of the regular and high level browsers have body weights nearly equivalent to that estimated for G. punjabiensis by Barry et al. (2002). According to Janis (1988), the HI of fossil species should be compared to extant taxa of similar body size/mass. The large giraffids; Sivatherium giganteum and Hydaspitherium-Bramatherium lineages exhibited dietary preferences of mixed feeders in open habitats (MFO) as are their extant analogues Loxodonta africana and Camelus dromedarius respectively. Lineages of Giraffa priscilla, G. punjabiensis, G. sivalensis were high level browsers (HB) as their HI is concordant with that of extant giraffids G. camelopardalis and O. johnstoni (okapi). HI and Body mass/size of Progiraffa exigua, Paleomeryx sp. are also consistent with that of a living Gazella Littocranius walleri and may be characterized as high-level browsers (HB) (Table 4.11; Figure 4.35; Appendix 12). 4.4.5. Tragulids: As regards the taxa of tragulids, Dorcath. majus, Dorcabune anthracotheroides, Dorcath. nagrii, Dorcatherium “259 sp.” (14.2-10.3), Dorcatherium 62
“311 sp.”(specimens of the studied taxa were recovered from fossil localities Y259 and Y311 respectively), Dorcabune nagrii were selective browsers in moist forested habitats. The Dorcatheriurn Y373 sp. was hypsodont (high crowned), and its succeeding taxon "Tragulidae/L101 sp." was even more hypsodont (Barry et al., 2002), signifying a change in dietary adaptations from browser/fruigivorous in closed, mixed feeder in semi-closed to grazer in open habitat. Archeotragulus (the ancestor of tragulids) inhabited in stable and warm forest in Pakistan by 18.0 Ma (Rossner, 2007) (Table 4.12). 4.4.6.Cervidae: Two species of large cervids Rucervus simplicidens and Cervus punjabiensis were regular browsers (BB) as Diceros bicornis and Diceros sumatrensis are (Appendix 12). Cervus sivalensis, C. rewati and C. triplidens and were mixed feeders in open habitat like Taurotragus oryx, Camelus bacterianus and C. dromedarius (Table 4.13; Figure 4.37). 4.4.7. Suidae: Potamochoerus “Y553 species”, Hippohyus sp., Sivahyus punjabiensis and Sus D013 species was browse dominated (HB) omnivorous in open woodland consistent with its extant species. Microstonyx major, P. hysudricus, Hippopotamodon sp. “Y450” H. sivalense, Tetraconodon magnus were fruigivorous/browsers in semi-closed woodland and resembled with B. babyrousa, P. porcus and Hylo. meinertzhazeni (Appendix 12). Lophochoerus nagrii was a small sized browser. The Conohyus sindiensis and L. pentapotamie were medium sized regular browsers (BB). Sivachoerus giganteus was a large plant dominated omnivorous animal. Palaeo. pascoei and H. chisholmi are known to have diverse dietary adaptations from browsers to fruigivorous. Bunolistriodon sp. was also frugivorous/browser (Table 4.14; Figure 4.38). 4.4.8. Equidae: Sivalhippus nagriensis is considered as regular browser (BB). S. theobaldi and S. perimense are presumed to be browsing mixed feeders. Equus sivalensis and Cremo. antelopinum may be placed among mixed feeders in open habitats. Sival. anwari and Equus sp. are inferred to be dry grass grazers (GG) (Table 4.15; Figure 4.39). 4.4.9. Rhinocerotidae: Alicornops aff. laogouense was regular browser (BB) whereas the dietary preferences of Alicornops complanatum changed from mixed feeder in closed habitats to open ones. The long lasting lineage of Chilotherium intermedium reflected the change in its dietary adaptation from regular browsers (BB) to mixed feeder 63
in closed habitat (MFC) and thereafter to MFO. Brachypo. perimense (SB/BB) and B. fatehjagense (B-W/SB) showed diverse dietary preferences ranging from browsers, selective browsers to mixed feeders in moist and swampy woodlands (amphibious- swampy, marsh and lake dewellers). Aceratherium sp. (BB) and Acerath.blanfordi (BB) were browsers and mixed feeders in open woodlands and grasslands. Rhinoceros sondaicus and R. sivalensis were mixed feeders in open habitats (MFO), and portrayed the dietary adaptations consistent with Camelus dromedarius and Loxo. africana, Gaindath. vidali and G. browni were regular browsers (BB) in woodlands. Punjab. platyrhinus may be considered as mixed feeder in open habitat (MFO). Hispano. matritense was found to be a small sized brachyodont-mixed feeder. Didermoceros aff. sumatrensis, Diderm. aff. abeli and Caementodon oettigenae were regular browsers (BB) (Table 4.16; Figure 4.40). The diet and habitat inferences of extinct species belonging to Anthracotheridae, Chalicotherini, Hippopotamidae and Proboscidea from the Pakistan Siwaliks are listed in appendix 5. 4.5. MEAN HYPSODONTY ANALYSIS Gradually increasing mean ordinated hypsodonty estimations in late Miocene shows seasonal climate with intensification of Asian Monsoon System (8.5-6.5) followed by progression of Modern Monsoon System during Plio-Pleistocene (Table 4.17) (see Table 5.1 and section 5.1 for details). During early Miocene, mean ordinated hypsodonty values show the warm and humid conditions with mean annual precipitation (MAP) 1500+ and mean annual temperature > 27C (Table 5.1; Figures 4.41, 4.42). During early middle Miocene (14-12.5 Ma) of Siwaliks, mean ordinated hypsodonty estimations (1.0- 1.4) suggest that the range of mean annual precipitation was 1500-1200 mm and annual temperature range was 22-26C. Hypsodonty values during late middle Miocene reflect the range of mean annual precipitation 1200-800 mm with annual temperature 20-24C (Table 5.1; Figures 4.41, 4.42). Hypsodonty analysis of early Late Miocene (11-9 Ma) localities shows that the mean annual precipitation was 1000-700 mm annual temperature range was 18-22C (Table 5.1; Figures 4.41, 4.42). Hypsodonty values (1.8-2.2) during the 9-8 Ma interval suggest the MAP was 800-600 mm and mean annual temperature was 15-18C. The annual precipitation patterns for later Late Miocene are different from earlier intervals that range from 700-500 mm with annual temperature 11-15C. During 64
the Pliocene (5-3.5 Ma), the mean hypsodonty estimations (2.5-2.7) reveal the range of mean annual precipitations 600-400 mm and annual temperature 10-12C. The precipitation patterns during late Pliocene (3.5-2.5 Ma) were ranging from 400-250 mm with annual temperature 8-10C. The 2.5-0.5 Ma interval shows hypsodonty values 2.7- 3.1 which indicate range of MAP 250-150 mm and mean annual temperature 6-8C (Table 5.1; Figures 4.41, 4.42). TABLES Table 4.1; Absolute and relative mesowear scorings of upper M2s of Giraffokeryx punjabiensis (Pilgrim, 1916). Abbreviations, n= number of samples. Mesowear Counts Percentages Cusp relief Cusp shape Cusp relief Cusp shape n High Low Sharp Round Blunt %High %Low %Sharp %Round %Blunt 17 17 0 11 6 0 100 0 64.70 35.29 0 Table 4.2; Absolute and relative mesowear scorings of upper M2s of Tragoportax sp. (Pilgrim, 1937). Abbreviations, n= number of samples Mesowear Counts Percentages Cusp relief Cusp shape Cusp relief Cusp shape n High Low Sharp Round Blunt %High %Low %Sharp %Round %Blunt 7 7 0 5 2 0 100 0 71.42 28.57 0 Table 4.3; Absolute and relative mesowear scorings of upper M2s of Selenoportax sp. (Pilgrim, 1937). Abbreviations, n= number of samples Mesowear Counts Percentages Cusp relief Cusp shape Cusp relief Cusp shape n High Low Sharp Round Blunt %High %Low %Sharp %Round %Blunt 12 11 1 8 4 0 91.6 8.33 66.66 33.33 0 Table 4.4; Absolute and relative mesowear scorings of upper M2s of Gazella lydekkeri (Pilgrim, 1937). Abbreviations, n= number of samples Mesowear Counts Percentages Cusp relief Cusp shape Cusp relief Cusp shape n High Low Sharp Round Blunt %High %Low %Sharp %Round %Blunt 12 8 4 7 5 0 66.6 33.3 58.33 41.66 0 65
Table 4.5; Data for Mesowear II and III for Extant Ruminants Species N Days Mesowear Mesowear Anterior Posterior Junction II III band 2 band 2 point "j" Browsers Okapia 11 0.8 1.4 1.5 1.3 1.5 johnstoni Giraffa 15 1.3 1.3 1.3 1.6 1.6 camelopardalis Alces alces 13 1.1 1.2 1.2 1.3 1.6 Grazers Ourebia ourebi 7 2.2 3.5 3.6 3.4 3.8 Kobus 10 3 3.1 3.1 3.1 3.5 ellipsiprymnus Chonnochaetes 12 4 3.8 3.8 3.8 3.7 taurinus Mixed feeders Cervus 4 1 1.6 1.6 1.6 1.5 Canadensis Grazella granti 17 2.2 2.5 2.3 2.6 2.9 First experiment Capra hircus 5 18- 1.8 1.7 1.3 2.2 2.5 Browsing 20 Capra hircus 5 18- 1.9 2.6 2.5 2.8 2.8 20 Second experiment Capra hircus 1 10 1.5 1.5 1 2 1 Browsing Capra hircus 1 20 1.5 1.5 1 2 1 Browsing Capra hircus 1 30 1.5 1.5 1 2 1 Browsing Capra hircus 1 40 1 1 1 1 1 Browsing Capra hircus 1 10 0 2 2 2 1 Grazing Capra hircus 1 20 1 2.5 2 3 2 Grazing Capra hircus 1 30 1 3 3 3 2 Grazing Capra hircus 1 40 1.5 3 3 3 2 Grazing The specimens of extant ruminants were used from the American Museum of Natural History.
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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 Fossil species from Mesowear Mesowear Anterior Posterior Junction Siwaliks of Pakistan II III band2 band2 point “J” Giraffokeryx 0.80 1.20 1.2 1.2 1 punjabiensis Pachyportax nagrii 2 1.87 2 1.75 0.75 E. khauristanensis 1 1.25 1 1.5 1 Gazella lydekkeri 1.75 2.16 2 2.33 1.66 Tragoportax sp. 1 1.34 1.43 1.37 1 Dorcatherium majus 1.08 1.58 1.5 1.66 1.16 Merycopotamus nanus 1.5 1.5 1.5 1.5 1.5 Miotragocerus gluten 2 2 2 2 2 Pachyportax latidens 2 3 3 3 3 Selenoportax lydekkeri 2.75 2.75 2.5 3 2.5 Selenoportax 3.25 2.87 2.75 3 2.5 vexillarius Hipparion sp. 4.9 3.85 3.85 3.85 3.78 Fossils of giraffids from China Samotherium boissieri 1.875 2.107 2.21 2.107 2.21 Samotherium sp. - 3 2 4 3 Samotherium sinense 3.33 3.33 3.33 2.6 Honanotherium 1.92 2.17 2.35 2 2.64 schlosseri Palaoetragus 2.11 2.66 2.66 2.66 2.86 coelophrys Paleotragus rouenii 1.75 2.5 2 3 2 Schasitherium tafeli 1.91 2.16 2.55 2.22 2.27 Specimens of Late Miocene giraffids from China used in this study are housed in the Paleontological Institute of Uppsala (PIU), Institute of Vertebrate Paleontology and Paleoanthropology at Beijing (IVPP), and the Hezheng Paleozoological Museum.
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Table 4.7; Fossil species from Late Miocene of Pakistan Siwaliks and their dental microwear results. Taxon N AP AS % LP % G % F % C % M %0-17
Giraffokeryx sp. 2 37.2 20.8 ------
Dorcatherium majus 36 26 17.43 81.3 28.1 15.6 3.1 81.3 50%
P.hysudricus 47 28.5 18.6 97.6 61 9.8 7.3 82.9 31.9%
H.sivalense 24 27.4 19.1 73.9 39.1 4.3 13 82.6 37.5% Selenoportax 6 17.41 17.58 ------vexillarius Tragoportax sp. 6 17 22.6 ------
Pachyportax latidens 4 18.8 28.37 ------
Miotragocerus gluten 1 12.5 18 ------
Merycopotamus 2 12 21.25 ------nanus Brach.perimense 1 11 22 ------
Hipparion sp. 1 9 27 ------
Brachy.fatejhangense 1 32.5 11 ------
Abbreviations: N, number of specimens; AP, average number of pits; AS, average number of scratches; %LP, percentage of individuals per taxon with more than four large pits, %G, percentage of individuals per taxon with gouges, %F, percentage of individuals per taxon with fine scratches, %C, percentage of individuals per taxon with coarse scratches, %M, percentage of individuals per taxon with a mixture of fine and coarse scratches.
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Table 4.8; Ecomorphic data of boselaphines from Siwaliks of Pakistan
Taxon N HI B.W (Kg) Diet AI Bovidae Bovinae Boselaphini Selenoportax sp. 04 2.70 50-250 MFC C Selenoportax falconeri - - 50-250 BB R Selenoportax giganteus 07 2.95 250-1000 MFO C Selenoportax vexillarius 06 1.50 50-250 BB C Pachyportax latidens 06 2.10 >250 MFO C Tragoportax sp. older 03 1.20 20-100 BB C Tragoportax sp. younger 04 1.89 20-100 MFC C Tragoportax salmontanus - - 50-250 BB - Kubanotragus skolovi - - 50-250 BB R Protragocerus gluten - - 50-250 BB R Miotrag. salmontanus small sp. 05 1.33 <15 SB C Miotragocerus gluten older 04 1.50 50-250 MFC C Miotragocerus gluten younger 05 1.60 50-250 MFO C Miotragocerus large sp. 04 2.10 >250 MFO R Helicoportax praecox 04 1.13 <15 SB C Helicoportax tragelaphoides - - 50-250 SB C Helicoportax sp. - - <15 SB R Elachistoceras khauristanensis 06 1.33 <15 SB C Ruticeros pugio - - >250 MFO C Tragoceridus sp. - - 50-250 MFO C Sivoreas eremite - - 50-250 MFC R Eotragus noyei 05 1.50 1-20 HB R Strepsiportax sp. - - - HB R Paleohypsodontus zinensis 04 2.28 - MFC R “2-Keel” a large bovid sp. - - 400-1000 MFO R Key to Symbols; “HB”= high level browser; “BB”= regular/unspecialized browser; “SB”= selective browser; “O”= omnivore “MFC”= mixed feeder in closed habitat; “MFO”= mixed feeder in open habitat; “FG”= fresh grass grazers; “GG”= dry grass grazer; AI=abundance index; C=common; A=abundant; R= rare. Estimated body weights/masses are from Badgley and Behrensmeyer (1980); Vrba (1980); Flynn et al. (1995); Nelson (2003); Brugal and Croitor, (2007).
69
Table 4.9; Ecomorphic data of antilopines of Siwaliks. Taxon N HI B.W (Kg) Diet AI Antilopini Antilope subtorta 05 2.08 125-343 MFO R Antilope planicornis 06 2.07 125-343 MFO C Antilope intermedia sp. nov 04 2.80 <343 GG C Antilope cervicapra - 20-100 GG R Gazella lydekkeri younger 06 1.98 >100 MFO C Gazella sp. older 05 1.65 20-100 MFC C cf. Prostrepsiceros vinayaki - - 20-100 MFC R Protragelaphus skouzesi - - 20-100 MFC R Hippotragini Hippotragini Y453 species - - >343 GG R Hippotragus brevicornis - - 125-343 GG C Hippotragus bohlini - - 280 GG C Oryx sivalensis - - 125-343 GG R Reduncini - - Reduncini/D013 species - - >343 FG C Sivacobus palaindicus - - 125-343 FG C Vishnocobus patulicornis - - 125-343 FG C Hydaspicobus auritus - - >343 FG - Sivadenota sepulta - - 125-343 FG C Indoredunca sterilis - - >343 FG R Kobus porrecticornis - - >343 FG - For the key to symbols of diet of fossil species see captions mentioned below the table 4.8. Estimated body weights were taken from Vrba (1980). Flynn et al. (1995); Brugal and Croiter, (2007) Table 4.10; Ecomorphic data of bovines from Siwaliks of Pakistan
Taxon N HI B.W (Kg) Diet AI Bovini Proamphibos lacrymans 05 2.56 >343 MFO C Proamphibos sp. 04 4.53 450 FG C Proamphibos kashmiricus - - - MFO - Hemibos triquetricornis 06 2.59 >343 MFO C Hemibos antilopinus - - >343 MFO - Hemibos acuticornis - - >343 MFO - Babalus palaindicus 05 2.85 465 FG C Bos sp. - - >343 MFO R Bos acutiferons 04 2.53 >343 MFO C Bison sivalensis 05 2.67 500 MFO R Bison crossicornis - - >343 MFO R Leptobos falconeri - - 320 MFO R For the key to symbols of diet of fossil species see caption mentioned below the table 4.8. Estimated body weights are from Badgley and Behrensmeyer (1980); Vrba (1980); Flynn et al. (1995); Nelson (2003), Brugal and Croitor, (2007). 70
Table 4.11; Ecomorphic data of giraffids from Siwaliks of Pakistan.
Taxon N HI B.W (Kg) Diet AI
Giraffidae Sivatheriinae Sivatherium giganteum 06 2.56 >500 MFO C Hydaspitherium megacephalum 05 2.00 250-1000 MFO C Hydaspitherium magnum 04 2.10 900 MFO C Giraffokeryx punjabiensis 06 1.31 233 MFC C Helladotherium grandi - - - MFO - Giraffinae Giraffa sivalensis 05 1.43 60-100 HB C Giraffa priscilla 06 1.33 - HB C Giraffa punjabiensis 04 1.32 60-100 HB C Progiraffinae Paleomeryx sp. 04 1.30 - HB R Progiraffa sp. - - HB C For the key to symbols of diet of fossil species see captions mentioned below the table 4.8. Estimated body weights are from Vrba (1980), Flynn et al. (1995); Barry et al. (2002) Table 4.12; Ecomorphic data of Siwalik tragulids. Taxon N HI B.W (Kg) Diet AI Tragulidae Dorcatherium majus 07 1.35 8-50 SB A Dorcatherium“270 species” - - <15 SB Dorcatherium nagrii - - <15 SB A Dorcatherium cf. majus - - 8-50 SB C Dorcatherium“259 species” small - - 8-50 SB C Dorcatherium sp. very small - - <15 SB R Dorcatherium“311species” - - - SB A Dorcatherium “373 species” - - - SB A Dorcatherium “457 species” - - - SB R Dorcabune anthracotheroides small 05 1.23 8-50 SB A Dorcabune nagrii - - 50-250 SB C Tragulidae /L101 unnamed sp. - - - FG R Archeotragulus - - <15 SB R For the key to symbols of diet of fossil species see caption mentioned below the table 4.8. Estimated body weights are from Vrba (1980). Flynn et al. (1995); Nelson, (2003)
71
Table 4.13; Ecomorphic data of cervids from Siwaliks of pakistan Taxon N HI B.W (Kg) Diet AI
Cervidae Cervinae Cervus sivalensis 06 1.80 125-343 MFO C Cervus triplidens 05 1.75 125-343 MFO C Rucervus simplicidens 04 1.37 <343 BB C Cervus rewati 05 1.87 125-343 MFO C Cervus punjabiensis 05 1.40 350 BB C For the key to symbols of diet of fossil species see caption mentioned below the table 4.8. Estimated body weights are from Vrba (1980). Table 4.14; Ecomorphic data of suids from Siwaliks of Pakistan.
Taxon N HI B.W (Kg) Diet AI Tayassuidae Pecarichoerus orientalis - - BB R Suidae Suinae Potamochoerus theobaldi 04 1.26 30-60 O R Potamochoerus palaindicus - - 8-50 O - Sus pregrinus - - 30-60 O C Hippohyus sp. 06 1.24 30-60 O C Sus D013 species 05 1.30 30-60 O C Sivahyus punjabiensis 04 1.17 - O C Microstonyx major 05 1.18 - O C Propotamoch. hysudricus 07 1.10 50 SB A Hippopotamodon “Y450 sp.” 04 1.13 50-250 O A Hippopotamodon sivalense 05 1.30 510 SB A Lophochoerinae Lophochoerus nagrii 04 1.20 - O R Conohyus sindiensis 06 1.12 8-50 BB A Tetraconodon magnus - 510 SB R Sivachoerus giganteus 04 1.15 510 O C Listriodontinae Listriodon pentapotamie 06 1.15 50-250 BB A Bunolistriodon sp. 05 1.10 - O R Hyotheriinae Hyotherium chisholmi 05 1.12 - BB A Paleochoeriinae Palaeochoerus pascoei 03 1.08 - BB R Palaeochoerus perimense - - - BB R For the key to symbols of diet of fossil species see captions mentioned below the table 4.8. Estimated body weights were taken from Vrba (1980). Flynn et al. (1995); Nelson (2003)
72
Table 4.15; Ecomorphic data of Siwalik horses. Taxon N HI B.W (Kg) Diet AI Equidae Equus sivalensis 05 3.11 350 MFO A Equus sp. 06 5.73 350 GG A Sivalhippus nagriensis - - 100-350 BB C Sivalhippus theobaldi 07 2.05 100-350 MFC C Sivalhippus perimense 06 2.10 100-350 MFC C Cremohipparion antelopinum 05 2.80 100-350 MFO A Sivalhippus anwari sp. nov 05 3.43 100-350 GG C For the key to symbols of diet of fossil species see captions mentioned below the table 4.8. Estimated body weights are from Vrba (1980); Flynn et al. (1995); Brugal and Croiter, (2007). Table 4.16; Ecomorphic data of Siwalik rhinoceroses Taxon N HI B.W (Kg) Diet AI Rhinocerotidae Alicornopini Alicornops aff. laogouense small sized 05 2.00 250-1000 BB R Alicornops complanatum - - >1000 MFO R Aceratherini Aceratherium sp. - - 1080 BB R Aceratherium blanfordi - - <1000 BB R Teleoceratini Chilotherium intermedium 05 1.33 1080- BB C Chilotherium intermedium 04 1.53 2700 MFC C Chilotherium intermedium 03 1.93 MFO C Didermoceros aff. sumatrensis - - <1000 BB - Didermoceros aff. abeli - - <1000 BB - Brachypotherium perimense 05 1.85 >1000 MFC C Brachypotherium fatehjangense 06 2.40 >1000 MFC R Rhinocerotini Rhinoceros sondaicus 05 2.50 1500 MFO C Rhinoceros sivalense 04 2.70 1500 MFO C Gaindatherium vidali 04 1.06 250-1000 BB R Gaindatherium browni 06 1.26 250-1000 BB C Punjabitherium platyrhinus 05 2.80 1500 MFO C Elasmotheriini Hispanotherium matritense 06 2.00 >1000 MFC C Caementodon oettigenae - - <950 MFC C For the key to dietary symbols of fossil species see captions mentioned below the table 4.8. Estimated body weights are gleaned from Vrba (1980); Flynn et al. (1995); Cerdeño, (1998); Cerdeño and Sánchez, (2000); Antoine, (2003); Brugal and Croiter, (2007).
73
Table 4.17; Mean hypsodonty values of major localities included in the study. Localities (age in Ma) Mean Hypsodonty values
Form (No. of taxa) SRDK-3 (73=ca. 1.2) 3.10 (3) SRDK-2 (642= ca.1.4 Ma) 3.00 (3)
Pinjor SRDK-1 (362= ca. 1.9-1.7) 3.00 (3) B0139 (3.3 Ma) 2.60 (4) B122 (ca.3.5 Ma) 2.55 (3) Kakrala (ca.3.5 Ma) 2.45 (3) Tatrot DKAWN (Rohtas fort) (ca.3.5 Ma) 2.55 (3) B044 (4.00 Ma) B119 (4.00 Ma) 2.57(3) 2.55 (4) B147 (4.50 Ma) B120 (4.50 Ma) 2.47 (3)2.40 (3) B151 (5.0 Ma) B152 (5.00 Ma) B150 (5.00 Ma) 2.55 (3), 2.50 (3), 2.40 (3) B157 (5.50 Ma) 2.45 (3) B0109 (6.20 Ma) 2.40 (4) B0110 (6.40 Ma) B0112, B0118 (6.40 Ma) 2.4 (5) 2.37 (3), 2.35 (3) B0135 (6.50 Ma), B0008, B0024 (6.50 Ma) 2.32.2.28, 2.20 B0007 (6.60 Ma) B0006 (6.60 Ma) 2.24 (4), 2.20(3) B0117 (7.20 Ma) 2.12 (5) B0104 (7.90 Ma) 2.15 (4) B0046 (8.0 Ma), B0043 (8.0 Ma) 2.18 (3), 1.97 (3) B0106 (8.10 Ma) B0103, B0025, 2.22 (5), 2.50 (3), 2.12 (5), B0026, B0027 2.25 (3), 2.15 (3). B0028, B0029, B0030 (8.10 Ma) 2.18 (3), 1.97 (2), 2.22 (4) B0031, B0032, B0033 (8.10 Ma) 2.20 (4), 2.15 (3), 2.10 (5) B0034, B0035, B0036 (8.10 Ma 2.05 (3), 2.0 (4), 2.0 (3) B0037, B0037 (8.10 Ma) 1.98 (4), 1.95 (6) B0040 (8.20) 1.98 (5) B0038 (8.30 Ma) B0105, B0044 (8.30 Ma) 2.00 (3), 1.98 (3), 1.97 (4) B0041, B0042, B0045 (8.30 Ma) 1.98 (4), 1.94 (3), 1.97 (3) Dhokpathan 74
Table 4.17 (continued)-Mean hypsodonty values of major localities included in the study Localities (age in Ma) Mean Hypsodonty values (No. of taxa) B0115 (8.80 Ma), B0111 (8.80 Ma) 1.90 (3), 1.87 (3) B09-13, (10 Ma) B017-23, (10 Ma) 1.82 (5), 1.93 (5) B107 (ca. 10.5 Ma) 1.79 (2) B108 (11Ma), B137, B155, B 159 (11 Ma) 1.59 (5), 1.57 (3),1.58 (4), 158 (3) Nagri B0123 (11.9- 11.7) 1.43 (2) B0054 (12.25 Ma) 1.39 (4) B0057 (12.70 Ma), B0055 (12.70 Ma 1.37 (4), 1.33 (3) B0056 (13.0 Ma) 1.29 (4) B0125 (13.40 Ma) 1.33 (2) B0051 (13.75 Ma) 1.36 (5)
Chinji B0048 (13.81 Ma) 1.38 (4) B0049 (13.81 Ma) B0050 (13.81 Ma) 1.33 (4), 1.37 (4) B0126 (13.83 Ma) B0047 (13.85 Ma) 1.28 (3), 1.36 (4) B0127 (14.0 Ma), B0052 (14.0 Ma) 1.38 (3), 1.28 (3) B0058 (14.21) 1.27 (3) Jaba, ca.15 Ma 1.06 (4)
DML ca. 16 Ma 0.93 (3) Kamlial Kamlial
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Table 4.18; Dietary characterization in extict giraffids based on p-values for the sample.
h with p-value with h with p-value with browsers browsers grazers grazers Giraffokeryx punjabiensis 0 0.801556617 1 0.000454653 Bramatherium small 0 0.613316 1 0.00354623 Bramatherium large 1 0.0372321 1 0.04675434 Pachyportax nagrii 0 0.7342167 1 0.00536781 E. khauristanensis 0 0.7654321 1 0.00432876 Gazella lydekkeri 1 0.0565343 1 0.00634562 Tragoportax sp. 0 0.6234521 1 0.00764321 Darcotherium majus 0 0.8123423 1 0.00865432 Dorcatherium minus 0 0.7432612 1 0.009543276 Merycopotamus nanus 0 0.6543721 1 0.00643251 Miotragoceros gluten 0 0.7543624 1 0.00432671 Pachyportax latidens 1 0.00745326 0 0.65423123 Selenoportax vexillarius 1 0.006342761 0 0.745321321 Hipparion sp. 1 0.005436721 0 0.854324145 Samotherium boissieri 1 0.048240422 1 0.003404859 Samotherium sinense 1 0.022118518 0 0.679128276 Alcicephalus neumayri 0 0.31865213 1 0.03541176 Honanotherium schlosseri 1 0.018803943 1 0.001675187 Palaoetragus coelophrys 1 0.00051262 1 0.012006729 Schasitherium tafeli 1 0.040085696 1 0.004347781 Palaeomeryx kaupi – artenay 0 0.145663019 1 4.42 E-16 Palaeomeryx kaupi – sanson 0 0.590057045 1 7.36 E-15
H = 0 accept null hypothesis (the samples are from populations with the same mean) p-value < 0.05
H= 1 reject null hypothesis (the samples are from populations with different means)
2-sided t-test assuming unequal variance.
76
100
80
60
40 Percentages 20
0 High Sharp Round Blunt relief cusp cusp cusp
Dental cusp relief and cusp shape Figure 4.1, Mesowear scorings displaying cusp relief (100% high) and cusp shape (64.70% sharp and 35. 29% round) of M2 of Giraffokeryx punjabiensis (Pilgrim 1916). No specimens with low cusp relief and blunt cusp shape were observed.
Number of ungulate species Figure 4.2, Dendrogram showing the hierarchical cluster analyses of Giraffokeryx punjabiensis amongst dataset of 27 extant species of ungulates comprising of browsers, mixed feeders and grazers from Fortelius and Solounias (2000). Abbreviation for the fossil species: Grf, punj= Giraffokeryx punjabiensis. Abbreviations for the extant species- Browsers: OH= Odocoileus hemionus, OV= Odocoileus virginanus, OJ= Okapia johnstoni, RS= Rhinoceros sondaicus, DS=Dicerorhnus sumatrensis, DB=Diceros bicornis, AA=Alces alces, GC= Giraffa camelopardalis. Grazers: ab= Alcelaphus buselaphus, bb= Bison bison, cs= Ceratotherium simum, ct= Connochaetes taurinus, dl= Damaliscus lunatus, eb= Equus burchelli, eg= Equus grevyi, he= Hippotragus equines, hn= Hippotragus niger; ke= Kobus ellipsiprymnus, rr= Redunca redunca. Mixed feeders: Me= Aapyceros melampus, Cc= Cervus Canadensis, Gg= Gazella granti, Gt= Gazelle thomsoni, Om= Ovibos moschatus, To= Taurotragus oryx, Ts= Tragelaphus scriptus, Ts= Tragilaphus strepsiceros. 77
A 2.5 2
HI 1.5 1 0.5 0 0 20406080100120 Occlusal relief
B 4 3 2 HI 1 0 0 20406080100120 Occlusal relief
C 8
6
4 HI
2
0 0 20406080100120 Occlusal relief
Figure 4.3 (A, B, C), Bivariate plots of hypsodonty index (HI) versus percentage of high occlusal relief; A= browsers, B= mixed feeders, C= grazers. Hypsodonty data were referred from Janis (1988), and mesowear data were taken from Fortelius and Solounias (2002) (●), Giraffokeryx punjabiensis (■) (Pilgrim, 1910). 78
A Browsers
6
4 HI 2
0 0 50 100 150 %age of sharp cusps
B Browsers 6
4 HI 2
0 0 20406080100120 %age of round cusps
C Browsers 2.5 2 1.5
HI 1 0.5 0 0 102030405060 %age of blunt cusps
Figure 4.4 (A, B, C), Mesowear analyses of cusp shape in G. punjabiensis (Pilgrim, 1910). G. punjabiensis (■) data for upper teeth are plotted against extant species of ungulates ●from Fortelius and Solounias (2000). In all bivariate plots, hypsodonty index is taken on y-axis and x-axis is represented by one of the three (% age of cusp shape) mesowear variables. 79
A Mixed feeders 4 3.5 3 2.5 2 HI 1.5 1 0.5 0 0 20406080100 %age of sharp cusps
B Mixed feeders 3.5 3 2.5 2
HI 1.5 1 0.5 0 0 50 100 150 %age of round cusps
C Mixed feeders 3.5 3 2.5 2 HI 1.5 1 0.5 0 012345 %age of blunt cusps
Figure 4.5 (A, B, C), Mesowear analyses of cusp shape in G. punjabiensis (Pilgrim, 1910). G. punjabiensis (■) data for upper teeth are plotted against the extant species of ungulates ●from Fortelius and Solounias (2000). In all bivariate plots, hypsodonty index is taken on y-axis and x-axis is represented by one of the three (% age of cusp shape) mesowear variables. 80
A Grazers 8 6 4 HI 2 0 0 20406080100 %age of sharp cusps
B Grazers 8 6
HI 4 2 0 0 20 40 60 80 100 120 % age of round cusps
C Grazers
8 6 4 HI 2 0 0 1020304050607080 %age of blunt cusps
Figure 4.6 (A, B, C): Mesowear analyses of cusp shape in G. punjabiensis (Pilgrim, 1910). G. punjabiensis (■) data for upper teeth are plotted against the extant species of ungulates ●from Fortelius and Solounias (2000). In all bivariate plots, hypsodonty index is taken on y-axis and x-axis is represented by one of the three (% age of cusp shape) mesowear variables. 81
120
100
80
60
Percentages 40
20
0 High sharp round blunt relief cusps cusps cusps
Dental cusp relief and cusp shape Figure 4.7, Mesowear scorings displaying cusp relief (100% high) and cusp shape (71.42% sharp and 28. 57% round) of M2 of Tragoportax sp. (Pilgrim, 1937). No specimens with low cusp relief and blunt cusp shape were observed.
Figure 4.8, Dendrogram showing the hierarchical cluster analyses of Tragoportax sp. amongst dataset of 27 extant species of ungulates comprising of browsers, mixed feeders and grazers from Fortelius and Solounias (2000). Abbreviations for the fossil species: Trg-sp= Tragoportax sp. Abbreviations for the extant species see captions for Figure 4.2. 82
A 2.5 2
1.5
HI 1 0.5
0 0 50 100 150 Occlusal relief
B 3.5 3 2.5 2 1.5HI 1 0.5 0 0 50 100 Occlusal relief
C 7 6 5 4
HI 3 2 1 0 0 50 100 150 Oclussal relief
Figure 4.9 (A, B, C), Bivariate plots of hypsodonty index (HI) versus percentage of high occlusal relief; A, browsers; B, mixed feeders; C, grazers. Data of living species were taken from Janis (1988), Fortelius and Solounias (2002) (●), Tragoportax sp. (■) (Pilgrim 1937). 83
100 90 80 70 60 50 40 Percentage 30 20 10 0 high relief low relief sharp cusps round blunt cusps cusps
Dental cusp relief and cusp shape Figure 4.10, Mesowear scorings displaying cusp relief (91.6% high), (8.33% low) and cusp shape (66.66 % sharp and 33.33% round) of M2 of Selenoportax sp. (Pilgrim 1937). No specimens with low cusp relief and blunt cusp shape were observed.
Figure 4.11, Dendrogram showing the hierarchical cluster analyses of Selenoportax sp. amongest dataset of 27 extant species of ungulates comprising of browsers, mixed feeders and grazers from Fortelius and Solounias (2000). Abbreviations for the fossil species: Selen. sp= Selenoportax sp. Abbreviations for the extant species see caption for Figure 4.2. 84
A 2.5 2 1.5
HI 1 0.5 0 0 50 100 150 Occlusal relief
B 4 3
HI 2 1 0 0 50 100 150 Occlusal relief
C 8 6 4 HI 2 0 0 50 100 150 Occlusal relief
Figure 4.12 (A, B, C), Bivariate plots of hypsodonty index (HI) versus percentage of high occlusal relief; A, browsers; B, mixed feeders; C, grazers. HI data from Janis (1988), and mesowear from Fortelius and Solounias (2000) (●), Selenoportax sp. (■) (Pilgrim 1937).
85
70
60
50
40
30 Percentage 20
10
0 high relief low relief sharp cusps round cusps blunt cusps
Dental cusp relief and cusp shape Figure 4.13, Mesowear scorings displaying cusp relief (66.66% high), (33.33% low) and cusp shape (58.33 % sharp and 41.66% round) of M2 of Gazella lydekkeri (Pilgrim 1937). No specimens with low cusp relief and blunt cusp shape were observed.
Figure 4.14, Dendrogram showing the hierarchical cluster analyses of Gazella lydekkeri amongst dataset of 27 typical extant species of ungulates comprising of browsers, mixed feeders and grazers from Fortelius and Solounias (2000). Abbreviations for the fossil species: Gz-ldkr= Gazella lydekkeri. Abbreviations for the extant species see caption for Figure 4.2. 86
A 2.5 2 1.5
HI 1 0.5 0 0 50 100 150 Occlusal relief
B 4 3
HI 2 1 0 0 50 100 150 Occlusal relief
C 7 6 5 4
HI 3 2 1 0 0 20 40 60 80 100 120 Occlusal relief
Figure 4.15 (A, B, C), Bivariate plots of hypsodonty index (HI) versus percentage of high occlusal relief; A, browsers; B, mixed feeders; C, grazers. Data from Janis (1988), Fortelius and Solounias (2002) (●), Gazella lydekkeri (■) (Pilgrim 1937). 87
Figure 4.16 (a), Rating/dietary characterization of browsers, mixed feeders and grazers on the basis of mesowear-III.
Figure 4.16 (b), shows the separation between the extant wild grazers and browsers using a 2 sample t-test assuming unequal variance. The difference between the two groups of animals was statistically significant. The mesowear of the mixed feeders overlapped with the mesowear of the grazers and the browsers. 88
4 3.5 3 2.5
Scale Grazing 2 Browsing 1.5 1 0 10203040 Time (days)
Figure 4.17, Results for the goat experiment. The green line shows a progression of the mesowear for the four goats from 10 day grazing to 40 day grazing. The teeth became incrementally flatter during the period of 30-40 days of browsing. The red line shows a progression of the mesowear for the four goats from 10 day browsing to 40 day browsing. The teeth became incrementally rounder for the first 30 days of grazing, and did not change during the last 10 days. At the starting point the pre-experiment goats had mesowear III range from 1.7-2.5.
Figure 4.18, Rating/dietary characterization of fossil species of giraffids from Siwaliks and from Late Miocene of China on the basis of mesowear-III. 89
Figure 4.19, Dendrogram is showing the hierarchical cluster analyses of 7 fossil species of boselaphines amongst dataset of 10 extant species of ungulates comprising of browsers, mixed feeders and grazers. Abbreviations for the fossil species: E.khaur= Elachestoceros khauristanensis, Trag.sp= Tragoportax sp., Pach. nag =Pachyportax nagrii, Pach. lat= Pachyportax latidens, M.gltn= Miotragoceros gluten, Selen.vex= Selenoportax vexillarius, Selen. lyd = Selenoportax lydekkeri. Abbreviations for the extant species- Browsers: OJ= Okapia johnstoni, GC= Giraffa camelopardalis, AA= Alces alces, CH= Capra hircus. Grazers: oo= Ourebia ourebi, ct= Connochaetes taurinus, ke= Kobus ellipsiprymnus, ch= Capra hircus, Mixed feeders: Cc= Cervus canadensis, Gg=Gazella granti. 90
Figure 4.20, Dendrogram is showing the hierarchical cluster analyses of Dorcatherium majus amongst dataset of 10 extant species of ungulates comprising of browsers, mixed feeders and grazers. Abbreviations for the fossil species: D. mjs= Dorcatherium majus. Abbreviations for the extant species see caption for Figure 4.19.
Figure 4.21, Dendrogram is showing the hierarchical cluster analysis of Giraffokeryx punjabiensis amongst dataset of 10 extant species of ungulates comprising of browsers, mixed feeders and grazers. Abbreviations for the fossil species: GP= Giraffokeryx punjabiensis. Abbreviations for the extant species see captions for Figure 4.19. 91
Figure 4.22, Dendrogram is showing the hierarchical cluster analysis of Hipparion sp. amongst dataset of 10 extant species of ungulates comprising of browsers, mixed feeders and grazers. Abbreviations for the fossil species: H.sp= Hipparion sp. Abbreviations for the extant species see captions for Figure 4.19.
Figure 4.23, Dendrogram is showing the hierarchical cluster analysis of Merycopotamus nanus amongst dataset of 10 extant species of ungulates comprising of browsers, mixed feeders and grazers. Abbreviations for the fossil species: Mry. Nanus= Merycopotamus nanus. Abbreviations for the extant species see caption for Figure 4.19. 92
Figure 4.24, Dendrogram is showing the hierarchical cluster analysis of Gazella lydekkeri amongst dataset of 10 extant species of ungulates comprising of browsers, mixed feeders and grazers. Abbreviations for the fossil species: Gaz. lyd= Gazella lydekkeri. Abbreviations for the extant species see caption for Figure 4.19.
93
Figure 4.25, Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and fossils suines/suids from late Miocene of Pakistan at 35X magnification. Convex hulls have been drawn around extant leaf browsing (marked with red color) (LB), fruit browsing (marked with green color) (FB), and extant grazing (marked with blue color) (GR) taxa for ease of comparison. Abbreviations for the fossil species: P. H= Propotamochoerus hysudricus; H. S= Hippopotamodon sivalense. Abbreviations for the extant species-Leaf browsers (LB): AM= Antilocapra americana, AA= Alces alces, DB= Diceros bicornis, CL= Camelus dromedaries, LW= Litocranius walleri, GC= Giraffa camelopardalis, TT=T. strepsiceros, BE= Tragelaphus euryceros. Fruit browsers (FB): fCG=C. niger, fCD= Cephalophus dorsalis, fCS= C. silvicultor, fCN= C. natalensis. Grazers (GR): bb= Bison bison, ab= Alcelaphus buselaphus, eg= Equus grevyi, ct= Connochaetes taurinus, hn= Hippotragus niger, eb= E. quagga; ke= Kobus ellipsiprymnus, Mixed feeders: Bt= Budorcas taxicolor, Ax= Axis axis, Cc= Cervus canadensis, Ca= Capricornis sumatraensis, Ci= Capra ibex, Cd= Cervus duvauceli, Cu= Cervus unicolor, Lg= Lama glama, Gg= Gazella granti, Lv= L. vicugna, Om= Ovibos moschatus, Oc= Ovis canadensis, Ti= Tragelaphus imberbis, Tq=Tetracerus quadricornis, To= Taurotragus oryx, Ts= Tragelaphus scriptus, Tr= Boselaphus tragocamelus (data on extant species from Solounias and Semprebon, 2002) (Appendix 11).
94
Figure 4.26, Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and fossil tragulids from late Miocene of Pakistan at 35X magnification. Convex hulls have been drawn around extant leaf browsing (marked with red color) (LB), fruit browsing (marked with green color) (FB), and extant grazing (marked with blue color) (GR) taxa for ease of comparison. Abbreviations for the fossil species: DT.MS=Dorcatherium majus; Abbreviations for the extant species see caption for Figure 4.25.
95
Figure 4.27, Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and fossil Boselaphines from late Miocene of Pakistan at 35X magnification. Convex hulls have been drawn around extant leaf browsing (marked with red color) (LB), fruit browsing (marked with green color) (FB), and extant grazing (marked with blue color) (GR) taxa for ease of comparison. Abbreviations for the fossil species: SL, VX= Selenoportax vexillarius; Pach.lat= Pachyportax latidens; Trag.sp= Tragoportax sp.; Mt.gltn= Miotragocerus gluten. Abbreviations for the extant species see caption for Figure 4.25.
96
Figure 4.28, Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and Giraffokeryx sp. from late Miocene of Pakistan at 35X magnification. Convex hulls have been drawn around extant leaf browsing (marked with red color) (LB), fruit browsing (marked with green color) (FB), and extant grazing (marked with blue color) (GR) taxa for ease of comparison. Abbreviations for the fossil species: Grf.sp= Giraffokeryx sp., Abbreviations for the extant species see captions for Figure 4.25.
97
Figure 4.29, Bivariate plot of the average number of pits versus average number of scratches in extant ungulatesand fossil species of anthracothroides from late Miocene of Pakistan at 35X magnification. Convex hulls have been drawn around extant leaf browsing (marked with red color) (LB), fruit browsing (marked with green color) (FB), and extant grazing (marked with blue color) (GR) taxa for ease of comparison. Abbreviations for the fossil species: MY.NS= Merycopotamus nanus, Abbreviations for the extant species see caption for Figure 4.25.
98
Figure 4.30; Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and fossil equids from late Miocene of Pakistan at35X magnification. Convex hulls have been drawn around extant leaf browsing (marked with red color) (LB), fruit browsing (marked with green color) (FB), and extant grazing (marked with blue color) (GR) taxa for ease of comparison. Abbreviations for the fossil species: H.SP= Hipparion sp.; Abbreviations for the extant species see caption for Figure 4.25.
99
Figure 4.31; Bivariate plot of the average number of pits versus average number of scratches in extant ungulates and fossils of Rhinos from late Miocene of Pakistan at 35 X magnification. Convex hulls have been drawn around extant leaf browsing (marked with red color) (LB), fruit browsing (marked with green color) (FB), and extant grazing (marked with blue color) (GR) taxa for ease of comparison. Abbreviations for the fossil species: BF= Brachypotherium fatejhangense, BP= Brachypotherium perimense; Abbreviations for the extant species see caption for Figure 4.25.
100
3.5 Helicoportax praecox Elachestoceros khauristanensis Tragoportax sp. older Miotragoceros salmontanus small sp. 3 Selenoportax vexillarius Eotragus noyei Miotragocerus gluten older Miotragocerus gluten younger Tragoportax sp. younger Pachportax latidens 2.5 Miotragocerus large sp. Paleohypsodontus zinensis Selenoportax sp. Selenoportax giganteus
2
1.5 Hypsdonty Index Hypsdonty
1
0.5
0 Boselaphine taxa
Figure 4.32; Hypsodonty analysis of boselaphine remains (HI=1.13-2.95). The results show the change in dietary and habitat adaptations from browsers/frugivorous to mixed feeders in closed and open habitats during 18.3-4.5 Ma.
Bovini 5
4.5 Bos acutiferons
4 Proamph.lycrym ans 3.5 Hemibos 3 triquetricornis Babalus 2.5 palaindicus Bison sivalensis 2 Hypsodoty Index Hypsodoty 1.5
1
0.5
0 Bovine species
Figure 4.33; Hypsodonty analysis of bovine remains (HI=2.56-4.53). The results show the change in dietary and habitat adaptations from mixed feeders (MFO) to grazers in open habitats during 14.2-0.5 Ma. 101
Antilopini
3
2.5
Gazella sp. 2 Gazella lydekkeri 1.5 Antilope planicornis Antilope subtorta 1 Antilope intermedia Hypsodonty Index Hypsodonty 0.5
0 species of antilopini remains
Figure 4.34; Hypsodonty analysis of fossils of antilopini (HI=1.65-2.80). The results show the change in dietary and habitat adaptations from mixed feeders in closed to grazers in open habitats during 14.2-0.5 Ma.
3 Sivatheriinae G. punjabiensis
2.5 Hydaspth. megacephalum 2 Hydaspth.magnum
Sivath. giganteum 1.5
1 Hypsodonty Index Hypsodonty
0.5
0 Sivatherine species
Figure 4.35; Hypsodonty analysis of sivatherine remains (HI=1.33-2.56). The results show the change in dietary and habitat adaptations from browsers (HB) to mixed feeders in open habitats during 14.2-2.5 Ma.
102
1.45 Giraffinae Paleomeryx sp.
1.4 Giraffa punjabiensis 1.35 Giraffa priscilla
1.3 Giraffa sivalensis
Hypsodonty Index Hypsodonty 1.25
1.2 Giraffine species
Figure 4.36; Hypsodonty analysis of fossils of Giraffinae (HI 1.3-1.43). The results indicate the persistence of high-level browsing adaptations till the Pleitocene of Siwaliks.
2 Cervidae
1.8 Rucervus simplicidens
1.6 Cervus punjabiensis
1.4 Cervus triplidens
1.2 Cervus sivalensis
1 Cervus rewati
0.8 Hypsodonty Index Hypsodonty 0.6
0.4
0.2
0 Fossil species of cervids
Figure 4.37; Hypsodonty analysis of fossils species of cervids (HI=1.37-1.87). The results show the progression of mixed feeders in open habitats at expense of browsers and mixed feeders in closed habitat during 5.3-1.5 Ma.
103
Suidae Palae. pascoei 1.4 Bunolistr. sp. L. pentapotamie 1.2 H. chisholmi
C. sindiensis 1 Hippopotam. sp.
0.8 Sivach.giganteus
P. hysudricus 0.6 Sivahyus punjabiensis
0.4 Microst. major
Hypsodonty Index Hypsodonty Sus D013 sp. 0.2 Hippohyus sp.
0 Potamoch. “Y553 sp” Taxa of suid remains Hippo. sivalense Figure 4.38; Hypsodonty analysis of fossil suids. The results show gradual increase in graze component at the expense of browse component in the diet during Siwalik succession (18.3-0.5 Ma).
Equidae Sivalh. theobaldi 7
6 Sivalh. perimense
5 Cremohip. antelopinum 4 Sivalh. anwari 3 Eq. sivalensis
Hypsodonty Index 2
1 Equus sp.
0 Fossil species of horses
Figure 4.39; Hypsodonty analysis of fossil equids (HI=2.05-5.73). The results show change in dietary and habitat adaptations from browsers to mixed feeders in closed and open habitats during 10.3-1.5 Ma.
104
Rhinocerotidae 3
Gaind. Vidali 2.5 Gaind. browni
2 Brachy. Perimense Chilo. intermedium 1.5 Hispano. matritense Alicorn. laogouense Hypsodonty Index Hypsodonty 1 Brachy. fatehjangense R. sondaicus 0.5 R. sivalensis Punjabi. platyrhinus 0 Fossil species of rhinoceroses
Figure 4.40; Hypsodonty analysis of fossils species of rhinoceroses (HI=1.1-2.7). The results reveal the progression of open habitat mixed feeders and grazers at expense of browsers and mixed feeders in closed habitats in the Siwaliks of Pakistan (18.3-0.5 Ma).
Figure 4.41; Map showing mean hypsodonty values for Siwaliks of Pakistan (16-0.5 Ma). 105
Figure 4.42; Map showing estimated precipitation ranges for Siwaliks of Pakistan (16-0.5 Ma).
106
CHAPTER 5
DISCUSSION The exploration of paleoenvironment of Pakistan Siwaliks has been largely based on faunal composition of mammals and functional aspects of their dental morphology such as hypsodonty (18.3-0.5 Ma), mesowear (18-5 Ma) and microwear (10.5-5 Ma) analyses. 5.1. PALEOCLIMATIC RECONSTRUCTION Reconstruction of paleoclimatic regimes of Siwaliks of Pakistan is based on progressive changes in mean hypsodonty values and species diversity (species richness and relative abundance) of mammalian fossil communities. The notable changes in mean hypsodonty values of ungulate fossil assemblages and in their species richness in association with ecological structure of living communities of mammals can explore climatic gradients at temporal and spatial resolution. The significant changes in mean ordinate hypsodonty values of assemblage of ungulate remains, their species richness together with ecological roles of extant communities are indicative of climatic gradients through times (Badgley and Fox, 2000; Jernvall and Fortelius, 2002). Ecomorphic data on ancient ungulates shows that the paleoclimate of Pakistan Siwaliks appears to change from hot and humid (planetary) to arid and seasonal or moonsonal one. The paleoclimate system of Neogene of Siwaliks is considered as the transition phase from hot and humid (green house) climate of paleogene to cold and arid (ice house) climate of Quaternary (Retallack, 2001; Zachos et al., 2001; Willis and McElvain, 2002; Bruch et al., 2007). The general trend of global climatic cooling was began in the Miocene epoch and continued through Pleistocene (Shakleton, 1995). Ecomorphic and mean ordinated hypsodonty data suggests that the early Miocene (18.3-16 Ma) of Siwaliks is predominated by brachydont browsers and forest frugivorous ungulate taxa which is indicative of humid climate with mean annual precipitation (MAP) of 1500+ mm and high (>27C) mean annual temperature (dense tropical and
multicanopied forest with C3 vegetation and marshy areas) (Tables 5.1, 5.3; Figures 4.41, 4.42). The early Miocene represents the general trend of climatic cooling that gradually progressed though Plio-Pleistocene as explored on the basis of mean values of 107
hypsodonty (Table 5.1). In habitats where mean annual rainfall is > 1500 mm and high mean annual temperatures, the type of climate is warm and humid (Janis et al., 2004). The 16-14 Ma interval showed increasing number of browsing and frugivorous species and mean hypsodonty values suggest the gradual declining of mean annual precipitation (1200-1500 mm) and mean annual temperature ranged from 22-26C (Table 5.1; Figures 4.41, 4.42). The evolving ecosystem showed less tropical canopy forest and
widening of tropical savannas with C3 grasses during this interval. The mammalian remains of early Miocene show slightest evidences of incipient aridity and the localities/sediments displaying increased hypsodonty values have been recorded from later part (16-14 Ma) (Fortelius et al., 2002). In the early Miocene of Pakistan Siwaliks, low mean hypsodonty values (0.93-1.26) show that the whole study area of Kamlial and early Chinji was relatively warm and humid (Table 5.1). In the 17–13.5 Ma (NMU5-6), low hypsodonty values show that the whole study area was warm and humid (Fortelius et al., 2002). Mesowear analysis of early Miocene taxa reflect the dominance of high and sharp dental cusp, mesowear scale ranging from 0-1, and sharp edges of dental enamel band two which indicates the warm and uniform paleoclimate (Appendix 2). During the Middle Miocene, the analysis of ecomorphic data on ungulates reveals the slight evidences of increasing aridity as the occurrence of ungulate assemblages are characterized by mixed feeding (MFC) bovids and hypsodont rhinoceroces (Hispanotherium lineages) but are still predominated by browsing ruminants, and suids. The mean hypsodonty values (1.2-1.8) and mean annual precipitation (MAP) values (800-1400 mm) with range of mean annual temperature (20- 24C) suggest the development of regionally and interregionally diversified climate system inconsistent with that of today’s patterns. Hence the middle Miocene climate of the Pakistani Siwaliks was relatively warm and humid, tropical or subtropical with initiation of Asian monsoon system (Table 5.1; Figures 4.41, 4.42). The hypsodonty based ecomorphic patterns before 11 Ma show the development of inter-regionally diversified paleoclimate system different to that of present days (Fortelius et al., 2002). During middle Miocene, mesowear signatures exhibit significant number of round and high cusps with abundance of sharp and high cusps (e.g., G. punjabiensis) which reveals the change in paleoclimate from warm and humid to tropical /subtropical one with the 108
initiation of paleoseasonality and birthing of South Asian monsoon system (Appendix 2, 7).The middle Miocene of Pakistan Siwaliks (Chinji-early Nagri Formations) is considered as the most critical interval “The Middle Miocene Transition” (Flower and Kennett, 1994; Zachos et al., 2001). Consequently, the latitudinal gradient increased through time and arid belts appeared in the middle latitudes (Flower and Kennett, 1994). The occurrence of continental mid-latitude drying in the later Neogene is well documented from Siwaliks including Europe, America, East Africa and Australia (Flower and Kennett, 1994; Fortelius et al., 2002; Barry et al., 2002; Ivanov et al., 2002; Böhme, 2003; Jiménez- Moreno and Suc, 2007). In particular, the contentious question of when the Asian monsoon became established complicates the issue. One view holds that the Asian monsoon began in the later Late Miocene, ca. 7–8 Ma (Sun et al., 1998; Ding et al., 1999, 2001; An et al., 2001), another view has it beginning much earlier, in the early Miocene (Guo et al., 2002; Sun and Wang, 2005). At the end of the middle Miocene and the beginning of the Late Miocene there was a further climatic change towards an increase in temperature and a decrease in humidity (Cerdeño and Nieto, 1995). Mean ordinated hypsodonty values (1.2-1.4), mean annual precipitation values (600-1000 mm) and temperature ranges (18-22C) during early Late Miocene (11-9 Ma) suggest the existence of seasonal climate and South Asian monsoon system (Table 5.1; Figures 4.41, 4.42). Furthermore, the relative abundance of ungulates during this interval shows the decreasing proportions of browsers (BB, HB; 32.5%-26.23%), frugivores/selective browsers (35-30.95%), browsing mixed feeders (18.9-16.7%) and increasing percentages of mixed feeders (2.7-11.90%) in open habitats and appearance of grazers (2.38%) which also led to development of seasonality and initiation of Asian monsoon system (Table 5.3; Figure 5.9). The early Late Miocene estimation of low mean hypsodonty values from Dhok Pathan stratotype and Hasnot areas revealed the persistence of humid conditions as were experienced in the region of central Europe. These findings are not in consonance with that of Eronen et al. (2010) on Siwaliks of Pakistan. According to Eronen et al. (2010) “during early Late Miocene of Pakistan Siwaliks, the mean hypodonty values showed increase in aridity with mean annual precipitation of < 500 mm”. The estimation of precipitation pattern on Siwaliks by Eronen et al. (2010) have generalized interpretation. This study presents more precise 109
estimation of precipitation and climatic regime than the earlier one. However, further compilation of mean hypsodonty data from more fossil localities of this interval may reveal even more precise precipitation estimation than the present study. Late Miocene of northern Pakistan shows climatic continuum of temperate and humid conditions, with clear seasonal differences (Barry et al., 2002). In the Late Miocene record of Pakistan Siwaliks, seasonality of precipitation, as inferred from annual profiles of 18O in equid teeth, followed a monsoonal pattern with a long dry season each year (Nelson, 2005). Mean hypsodonty data (1.4-2.4), mean annual precipitation estimates of 800-600 mm and range of mean annula temperature (11-15C) during mid Late Miocene to latest Miocene (9-6 Ma) reveal that the existing of pronounced seasonality and intensifiation of South Asian monsoon system in this interval (Table 5.1; Figures 4.41, 4.42).The relative abundance of ungulates during this interval shows the decreasing proportions of browsers (BB, HB; 26.23-10.56%), frugivores/selective browsers (30.95-10.52%), browsing mixed feeders (16.7-2.63%) and increasing percentages of mixed feeders (11.9-44.74%) in open habitats and grazers (2.38-18.42%) which also led to development of pronounced seasonality and intensification of Asian monsoon system (Table 5.3; Figures 5.5, 5.9). Change in paleocommunity structure of Siwalik mammals indicates that species diversity and increased episodes of faunal turnovers in Late Miocene are concordant with global climatic regimes of increased aridity and paleoseasonality. During 10.0 to 6.3 Ma intervals, the 18O signatures of equid dental profiles shows a decline in mean annual rainfall by several hundred mm, while the paleoseasonality of precipitation remained constant. It is consistent with present day
ecotone in South Asia existed between savanna (C4) and dry deciduous /monsoon forest
(C3) (Badgley et al., 2008) which is maintained by small periodic changes in the duration of dry season, grazing pressure, or occasional fire (Stott, 1990; Mariotti and Peterschmitt, 1994). For the Late Miocene record, the vegetation change also exhibits the transformation of a three dimensional habitat to a more two dimensional one along with a decreased variety of trophic resources. These paleoclimatic and paleoecological changes were driven by the uplift of the Himalayas and Tibetan Plateau that, in turn, intensified the South Asian monsoon system (Zhisheng et al., 2001). Enrichment in 18O values from pedogenic/paleosole carbonates and mammalian dental profiles at 9 Ma represents 110
an elevation in temperature, and a decrease in paleoprecipitation, or a change in source of precipitation-any one of which may be synchronized with the onset of South Asian/Indian monsoon system (Quade and Cerling, 1995). Isotopic historical trends were documented for mammalian herbivores and for individual lineages over the Late Miocene interval of environmental change. Mammalian 13C values show a greater range of variation than do those from pedogenic carbonates. This variation reflects dietary differences among individuals and species as mammals ranged across the sub-Himalayan flood basin and documents the presence of more closed and more open habitats than are represented by paleosol carbonates (Nelson, 2007). Badgley et al. (2008) evaluated the isotopic signatures and ecomorphological dietary data for individual mammalian lineages in relation to emergences and demises of taxa to test the premise that climatically induced paleovegetation change which in turn triggered the observed mammalian faunal turnover. The great diversity of the animal community reflects the availability of diverse habitats and intensification of South Asian monsoon system during 8 to 7 Ma (Wang et al., 1999) leading to dry and more seasonal paleoclimate. The 18O values are indicative of changes in precipitation pertaining to a drier and more seasonal paleoclimate. By 7.3 Ma, 18O signatures typical of middle Miocene of Siwalik rocks are not evident (Quade and Cerling, 1995). It is admitted evidence that the decline in mammalian species diversity and increased faunal turnovers in Late Miocene of Siwaliks could be synchronized with global climatic inferences of elevated aridity and paleoseasonality (Barry et al., 1995). Mesowear analysis shows that the average mesowear-III scores in Late Miocene Siwalik taxa have been increasing from 1.20-3.85 with junction points (JPs) from 1.33- 3.78 through times (Appendix 2). Results of both mesowear-II and mesowear-III reveal a gradual increase in mesowear scorings with high and sharp cusps becoming less frequent and increased prevalence of round, high, low and blunt cusps (Appendix 2, 7). These progressive shifts in mesowear signatures indicate a trend toward a decreased browse component and an increased graze component in the diet of Siwalik ungulates while retaining their mixed feeding strategies. This trend of increasing low and blunt cusps reveals the progressive change in paleoseasonality throughout Late Miocene intensification of South Asian monsoon system during 8-6 Ma interval. Ever increasing the proportions of high with round, low and blunt cusps in latest Miocene taxa also 111
indicate the initiation of modern monsoon system. The geo-historical record of South Asia (ca. 8 and 6 Ma) observed by Badgley et al. (2008) and the mesowear study on hipparionine horses from Late Miocene (10.3-6 Ma) of Pakistan Siwaliks worked out by Wolf et al. (2013) and also support the findings of this new mesowear study. This paleoenvironmental evolution has been concordant with increased upheavals of the Himalayas that triggered the changes in precipitation patterns due to the onset of the Indian monsoon system (Quade and Cerling 1995; Zhisheng et al., 2001; Barry et al., 2002; Behrensmeyer et al., 2007). Microwear signatures of fossil species of two suines (Propotamochoerus hysudricus and Hippopotamodon sivalense), a tragulid (Dorcatherium majus) from Late Miocene of Siwaliks exhibit the presence of fine, coarse and mixed scratches with variable number of pits which indicate the essence of South Asian monsoon system with pronounced seasonality during early Late Miocene. The individual microwear assessment of the studied taxa shows the increasing proportion of variable number of pits with coarse and mixed scratches through Late Miocene. These results also suggest that paleoseasonality successively intensified during younger intervals of Late Miocene. Microwear study of an assemblage of large boselaphines, an anthracothere, two rhinoceroses and a hipparionine horse from latest Miocene (8-6 Ma) reveal the decreasing number of pits and increasing number of scratches through times which indicate the intensification of South Asian monsoon system. The existence of excessive number scratches during latest Miocene (7-6 Ma) also suggest the initiation of modern monsoon system at 5 Ma (Appendix 3). This study also testifies the hypothesis that the most of the ungulate taxa from Siwaliks, who did not changed their dietary and habitat adaptations, went extinct in the order of their susceptibility. Nelson (2003) worked out the microwear study of fossil species of suines, Sivapithecus, tragulids, large and small bovids, giraffids, anthracotheres, gomphotheres, rhinos and hipparionine horses from Late Miocene Siwaliks of Pakistan. Paleoclimatic inferences drawn in her study are in consonance with the findings of the present study. Mean hypsodonty data (2.5-2.7) shows that the early Pliocene (ca. 5.5-4 Ma) of Pakistan Siwaliks received mean annual rainfall between 500 and 700 mm accompanied by mean annual temperature 10-12C (Table 5.1; Figures 4.41, 4.42). The relative 112
abundance among ungulates during this interval chronicles the decreasing proportion of browsers (BB, SB, HB; 21% to 18%), increasing proportion of mixed feeders in open habitat (MFO; 44.74% to 48.71%) and that of grazers (FG, GG; 18.42-23%) (Table. 5.3; Figures 5.5, 5.9). Integrated analyses of the data suggest the the birthing of modern monsoon system occurred in this interval. Patnaik (2011) proposed that the modern monsoon system we experience now days might have initiated by 5 Ma. This is supported by present day upheaval of the Tibetan Plateau that was attained at early Pliocene (Prell and Kutzbach, 1992). The distribution and relative abundance of Pliocene ungulates and murids overlap clearly with that of the modern monsoon region (surface based) (Patnaik, 2003). Furthermore, almost all ungulate and murid genera from Pliocene deposits in Siwaliks have living relatives inhabiting in the monsoon region (Downing et al., 1998; Van der Meulen and Musser, 1999; Patnaik, 2000, 2003). During the Late Pliocene (4-2.5 Ma) the mean hypsodonty ratios (2.5-2.8) revealed the precipitation values were ranging from 500 and 300 mm with range of mean annual temperature 8-10C (Table 5.1; Figures 4.41, 4.42). Moreover, the decreasing proportions of browsers (18-10.50%), stable occurrence of mixed feeders in open habitats (MFO; 47.5-48%) and increasing proportions of grazers (FG, GG; 23-36%) (Table 5.2, 5.3) shows more strengthening of modern monsoonal climate system. The Late Pliocene (4-2.5 Ma) precipitation values were ranging from 300 and 500 mm (Fortelius et al., 2006). Late Pliocene diversity of ungulate groups such as bovines, rudencini, antelopes, boselaphines, large giraffes, cervids, equus horses, rhinos and proboscideans with murine rodents reflects the existence of the diverse spectra of ecological niches, mainly open grassland/savanna conditions interspersed with dry/wet deciduous and semi-evergreen forests (Patnaik, 2011). Ecomorphic data of Pleistocene (2.5-0.5 Ma) ungulates of Siwaliks reveals the increasing proportions of mixed feeders in open habitats (MFO; 47-56%), with stable occurrence of grazers (35%) and browsers (10%) (Table 5.3; Figures 5.5, 5.9). Mean ordinated hypsodonty (2.8-3.0) also show the mean annual precipitation and temperature ranges of 200-300 mm and 6-8C respectively (Table 5.1, 5.3; Figures 4.41, 4.42). Combinig the results of both indicate the existence of arid and modern monsoonal climate with arid and cool winters, wet and warm summers during this interval. Terminal 113
Pleistocene around Pabbi Hill of Pakistan was relatively drier as compared to the Pliocene with decreasing mean annual rainfall upto 200 mm indicating the more aridity and intensification of modern monsoon system (Table 5.1). The Late Pliocene and Early Pleistocene sequence of the Pabbi Hills was also formed under a modern monsoonal system that intensified after the Late Miocene, with pronounced contrasts between cool and arid winters and warm and wet summers (Dennel et al., 2007). It has been observed that early Pleistocene (ca. 2.5 Ma) global climate change towards a drier, cooler and more variable one mainly linked with the glaciations processes of Northern Hemisphere (Mangerud et al., 1996; Williams et al., 1999). It appears that the diversification of ungulate and (published) murid faunas inhabiting in monsoonal sites indicate the modern monsoonal circulation that was intensified by Plio-Pleistocene upheavals of Himalayas and Tibetan Plateau. The diversified occurrence of murids has some how contributed to the monsoonal circulation that was greatly strengthened by uplift of the Himalayas during Plio-Pleistocene (Dewey et al., 1988). Data on body size of ungulate remains has been treated as paleoecological proxy for paleoclimatic variables (temperature ranges, precipitation patterns etc.) and paleohabitat inferences. Smaller body size of Siwalik ungulate lineages indicates warmer climate, closed-semiclosed or forested habitats whereas larger body size represents cooler and seasonal climate with open habitat such as savannas/grasslands (Tables 4.8-4.16). Analyses of ungulate body size distribution of Siwaliks succession of Pakistan suggest that smaller body sized forest dependent species evolved to larger and open grassland species which are consistent with the other paleo-climatic data (Table 5.1). Body size of ungulates has inverse relationship with mean annual precipitation and temperature ranges i.e. smaller the body size the higher will be the mean annual temperature, precipitation ranges and vice versa. Larger body size can prevent the escaping of heat across the body surface which is advantageous adaptation in colder climates while smaller body size can facilitate dissipation of heat that is beneficial in warm climate (Asthon et al., 2000). According to Badgley and Behrensmeyer (1980), Late Miocene faunas from the Potwar Siwaliks showed 15 over 29 ungulates <100 kg. This implies that the existence of diverse niches of vegetation accommodating the significant number of browsers with abundant cover. 114
Ungulate remains recovered from Pabbi Hills (Pinjor Formation) of Pakistan had not only the ranges of smaller body size (ca.1.7 Ma), but all the taxa (except Gazella and probably one type of suid) possessed body weights about 4200 kg (Dennel et al., 2006). The changes in body size of herbivorous mammals during the Late Miocene of Siwaliks can be coincided with more open habitats (Morgan et al., 1995). The increase in body size began with the first emergence of hipparionine horses’ ca.10.3 Ma (corrected
following Badgley et al., 2005) and the first appearance of C4 grasses at about 9.4 Ma (Morgan et al., 1994) and became more prevalent during 9-8.5 Ma. The progressively increasing body size of predators may also have attributed to the increased body size of mammalian herbivores, but it was treated as inconclusive premise. 5.1.1. IMPACT OF PALEOCLIMATE CHANGE ON FAUNAL EVOLUTION It is an admitted paleoecologic observation that paleoclimatic changes have been impacting on the evolution of ecosystems through times and Siwaliks of Pakistan is not exception in this regard. Ecomorphic data of ungulates suggest that the multiple episodes of paleoclimatic changes influenced the geographic ranges, abundance and morphology of individual taxa that in turn stimulated speciation, immigration, extinction events within mammalian paleocommunities though Siwalik succession. Integrated analyses of these processes punctuated the long-term patterns of faunal turnover, species diversity and paleocomminty stability in the Siwaliks of Pakistan. The climatic changes have been concordant with mammalian geographic range shifts, fragmentation and selective filtering of their populations over spatio-temporal scales leading to allopatric speciation, extinction and biotic turnover within the ecosystems and among continents (Rosenzweig, 1995; Vrba, 1995; Parmesan, 2006; Van Dam, 2006). The change in climatic shift regulated the change in paleovegetation from closed/C3 vegetation to semi-closed (C3/C4
transition) vegetation and then semi-closed to open vegetation system comprising of C4
grasses. Plant communities with predominantly C3 vegetation (closed vegetation) were greatly diminished after 7.0 Ma, and those with predominantly C4 vegetation (open vegetation), emerged as early as 7.4 Ma (Cerling et al., 1997; Barry et al., 2002). The successive changes in plant and herbivore paleocommunities observed during the latest Mio-Pleistocene are linked with generally accepted trends in climate change towards cooler, drier conditions, and increased seasonality (Patnaik, 2011). The climatically 115
induced changes in vegetation reveal that the climate triggers the change in vegetation habitats to expand, shrink, or migrate across the landscape, and herbivorous species dependant on particular vegetation or trophic resources will shift their biogeographic ranges to occupy the preferred habitats (Vrba, 1995). The general trend of global climatic cooling was began in the Miocene epoch and continued through Pleistocene (Shakleton, 1995) and its associated considerable aridity, increased seasonality may have contributed to the progression of more open landscapes (Rivals and Athanassiou, 2007). Warm and humid (planetary) paleoclimate of early Miocene (18.3-14 Ma) allowed to appear 2-4 browsing and fruigivorous taxa of suids, 2-6 fruigivorous lineages of tragulids, 2 lineages of giraffids, 2-8 browsing/fruigivorous lineages of boselaphines, 1-4 fruigivorous/browsing species of anthracotheres, four browsing and one fruigivorous species of proboscidea, 3 browsing, 3 mixed feeding and one fruigivorous lineages of rhinoceroses (Table 5.2). In the Middle Miocene of Siwalik record, this premise implies that warm and humid climate permitted to immigrate 8 browsing and fruigivorous taxa of suids, 4 fruigivorous lineages of tragulids, 3 lineages of giraffids, 5 mixed feeding and 4 fruigivorous lineages of boselaphines, one browsing species of antelopes, four fruigivorous/selective browsing and two mixed feeding species of anthracotheres, four browsing and one fruigivorous species of proboscideans, 2 browsing, 3 mixed feeding and one fruigivorous lineages of rhinoceroses. The subtropical paleoclimate during 12.5- 11.5 Ma reflect the few extinctions of browsing and mixed feeding lineages of rhinos and boselaphines and appearance of subtropical forest dependant taxa such as Gaindatherium browni, and Sivaceros gradiens. The moist and seasonal climate during 11.5- 10 Ma triggered the extinction of one frugivorous lineage of tragulids, two browsing lineages of giraffids and one species of suids while the appearance of seasonal and forest/woodland dependant two lineages of suines and 2 taxa of hipparionine horses. During 10-9 Ma, the temperate and seasonal climate reflect the demise of two (one MFC and other fruigivorous) lineages of boselaphines, one lineage of large suid; Tetraconodon magnus, one fruigivorous lineage of anthracotheres and appearance of two (one was browser and other was MFO) taxa of boselaphines, 3 (2 browsing, one fruigivorous) lineages of proboscidea and one lineage (Sus sp.) of suids. The intensification of South Asian 116
monsoon system caused shrinking of C3 vegetation and increasing mosaics of C3-C4 transitions reflect decreasing proportions of browsers, frugivores/selective browsers, browsing mixed feeders and increasing percentages of mixed feeders in open habitats and grazers during 9-8 Ma. The cascade of extinction events during Late Miocene of Siwaliks suggest that browsing, fruigivorous and closed habitat taxa together with their different thresholds for size of forest and fruit/browse abundance requirements went extinct at different rates following the same pattern of extinctions or responses of extant species to forest disturbance in modern ecosystems. The first group of species that went extinct was the closed and wet habitat species i.e. a large body sized fruigivorous ape and two taxa of tragulids. Subsequent extinctions were comprised of more fruit-dominated omnivorous suids, more taxa of tragulids with the most foliovorous species of tragulid under investigation were the last ones to go extinct. That gradient of extinction was most probably a response to fragmentation of forest and the species responded or demised at different time intervals because of their different susceptibilities (Nelson, 2003). Global temperatures dramatically declined during latest Miocene, as did taxonomic diversity of mammals, and the vegetational habitats at mid-latitudes became apparently more arid
(Fortelius et al., 2002). At ca. 8 Ma, a shift in C3-photosynthetic pathway to the C4 one happened in the lower latitudes in South Asia and North America (Cerling et al., 1993, 1997). The general pattern of evolution in the Late Miocene-Pliocene ungulates of Pakistan Siwaliks reveals that browsing lineages (tragulids and cervids) were replaced by grazing/mixed feeding in open habitats (bovids, equids etc.) rather than the brachydont browsers themselves under going change in their diet (Barry et al., 1995). Similarly evolved lineages of ungulate do not transform their dietary preferences back into browsers once they have changed the more derived masticating machinery of mixed feeders or grazers (Janis, 2008). During the interval, 8-6.5 Ma, dry and pronounced seasonal paleoclimate punctuated the genesis of semi-closed vegetation system that provided platform for great faunal transformation from closed habitat browsing and mixed feeding to that of open ones with progressive influx of grazers. The faunal turnover based on biostratigratigraphic data reveals that most of species reliant on C3 vegetation (fruit,
browse, and forest dependent) disappeared during the C3-C4 transition to C4 vegetation 117
(open grasslands), resulting in local extinction of these lineages, whereas immigrant
herbivores were grazing mixed feeders (C3-C4) or grazers (C4 vegetation). Thus, the pattern of faunal change might have involved local extinction of frugivores and browsers
and replacement by grazing mixed feeders and grazers. Consumers of C3 vegetation
could persist by changing their diets to include C4 vegetation. This premise is especially
plausible for ungulate lineages that consumed substantial quantities of C3 grasses before
the vegetation transition and then switched to C4 grasses. The significant number of long lasting lineages belonging to group hipparionine horses, rhinoceroses, boselaphines, sivatherines, antelopes and a tragulid altered their feeding preferences from browsers to mixed feeders in open habitats/grazers. However, more sampling and investigation is required to strengthen this premise. During the interval of greatest paleo-environmental
change (ca. 8.6 to 6.5 Ma), C4 vegetation replaced C3 vegetation and mammalian taxa
dependant on C3 vegetation may have declined in their abundance instead of having
disappeared locally, immigrants taxa incorporated C3-C4 (mixed feeders) and and C4-
diets (grazers). The sequencial move from C3-diet to C3-C4 to C4-diet reveals that savannas followed by grasslads replaced forests on the floodplain (Badgley et al., 2005). The evolutionary pattern of ungulates in the Late Miocene of Pakistan Siwaliks is indicative of gradual replacement of browsing and fruigivorous lineages by ones preffering more fibrous foods (bovids, equids etc.), rather than the browsers/fruigivores themselves undergoing dietary evolution. Similarly, lineages of ungulate do not transform dietary preference back into browsers once they have attained the more derived cranio- dental machinary of mixed feeders or grazers (Janis et al., 2008a, b).
These historical isotopic trends reveal that some species maintained C3-diets
while others changed their dietary habits incorporating varying amounts of C4 grasses
during the transitional phase from C3 to C4 grasses (Badgley et al., 2008). The 8.5 to 6.0 Ma intervals encompasse the 13C shift as well as the major duration of faunal turnover.
Prior to 8.5 Ma, most of the herbivores used to consume C3 grasses from closed forested
13 habitat ( C values -12‰). C3 vegetation was also a component of the flood plain cover. Mixed feeding and grazing lineages of artiodactyls and rodents showed a different turnover pattern revealing the disappearance of 11 lineages and appearance of 9 lineages during 8.5 and 6.0 Ma (Badgley et al., 2008). Four taxa showed enrichment of 13C 118
signatures in younger intervals, and 2 of the lineages persisted beyond the major faunal
turnover. One lineage of bovid used to feed on C3 diet at 8.0 Ma that changed to a C4- diet at 7.5 Ma afterwards. Hipparionine lineages incorporated some C4-diet by 8.5 Ma (Quade et al., 1994; Macfadden et al., 1999a, b). New taxa of mixed feeders and grazers appeared throughout this major interval contrary to the browsers and frugivores (Badgley et al., 2008). Overall ungulate species richness declined. As regards rodents, hypsodonty successively increased in one long-lasting lineage of rhizomyid whereas other hypsodont lineages of rhizomyid appeared after 7.5 Ma (Flynn and Jacobs, 1982). This pattern of mammalian faunal turnover shows prolonged paleoclimatic forcing on faunal evolution of Siwalik Miocene (Badgley et al., 2008). It has been found that this turnover event had a profound impact on the grazing species of mammals and a greater biodiversity of hypsodont forms of artiodactyls including the bovids emerged in Pakistan during this interval (ca. 8 Ma) whereas brachydont forms such as giraffids and suids declined (Barry
et al., 1995). The inhabitants of mixed (C3/C4) communities or the humid regions of the floodplain were presumed to have been extinct in increasingly open habitats during the latest Miocene and the pace of extinction events increased to 3/4th after the emergence
and dominance of C4 grasses on the floodplain (Barry et al., 2002). The transitional phase between the Miocene and the Pliocene faunas appears to be triggered by the Miocene climatic optimum (Figueirido et al., 2011). The development of modern monsoon system due to uplifting of Tibetan Plateau at 5 Ma regulated the complete extinction of one fruigivorous tragulid and appearance of two lineages of cervids among browsers, three lineages of cervids among closed habitat mixed feeders, two lineages of giraffids, one antelope and one boselaphine among mixed feeders in open habitats. The extinction events happened in accordance with the intensity of modern monsoon system and susceptibility of population. The Pliocene witnessed a successive and sharp increase in paleseasonality that triggered the faunal evolution displaying the savanna mosaic characters. Grazing hippos and antelopes replaced anthracotheres and tragulids as the dominant representatives of artiodactyls. As regards perissodactyls, browsing rhinos were replaced by grazing ones and hipparionines were replaced by three-toed equids. Hyosodont Elephas taxa replaced bunodont and long jawed gomphotheres. 119
Conversely at 3.5 Ma, the modern moosonal climatic regime magnified the influx of two lineages of proboscidea, 5 lineages of bovines in open habitat mixed feeders and 2 browse dominated omnivorous suids. As regards grazers, there was demise of hipparionine horses and appearance of two lineages of Equus horses and five bovines and two reduncini. The early Pliocene faunal diversity coincides with the warming during that interval and its subsequent decline synchronizes with later cooling process (Figueirido et al., 2011). The elevation of the Pliocene fauna can be linked with the major immigration events that happened at about the end of early Miocene (Janis, 1993). During 2.5-0.5 Ma, intense modern monsoonal climate caused the extinction of one suid, one grazing tragulid and influx of two giraffids and three lineages of grazing rhinoceroses. It appears that the diversification of ungulate fauna and published work of murids inhabiting in monsoonal sites indicate the modern monsoonal circulation that was intensified by Plio-Pleistocene upheaval of Himalayas. The diversified occurrence of murids has some how contributed to the monsoonal circulation that was greatly strengthened by uplift of the Himalayas during Plio-Pleistocene (Dewey et al., 1988). The late Pleistocene extinction events did involve a profound decline in species richness and immigration of modern man into all the continents. A loss of significant number of ungulates was due to synergistic effect of drier paleoclimate and hunting of humans (Koch and Barnosky, 2006). 5.2. EXPLORATION OF PALEOVEGETATION The ecomorphic study of Siwalik ungulate dentition, and faunal composition of the area suggest that the vegetation configuration of Siwaliks changed from closed
vegetation system (C3 vegetation) (18.3-8.5 Ma) to semi-closed one and from semi-
closed vegetation system (C3-C4 vegetation) (8.5-6.5 Ma) to an open vegetation system
(C4 vegetation) (6.5-0.5 Ma). Plant communities with predominantly C3 vegetation were
greatly diminished after 7.0 Ma, and those with predominantly C4 vegetation (open vegetation), emerged as early as 7.4 Ma (Cerling et al., 1997; Barry et al., 2002). Early Miocene (18.3-16 Ma) of Siwaliks is predominated by low species richness of browsers (5-11) and forest frugivorous (2-3) ungulate taxa which is indicative of dense
and multicanopied tropical rain forest (C3 vegetation) with limited mosaics of tropical savanna like habitats (Table 5.2). Late early Miocene (16-14 Ma) showed increasing 120
number of browsing (11-19) and frugivorous species (3-8) which suggest that the evolving ecosystem showed less tropical canopy forest and widening of tropical savannas with more prevalence of C3 grasses than the earlier interval (Table 5.2). As regards mesowear analysis, the early Miocene ungulate taxa shows the predominance of high and sharp dental cusp with mesowear scale (0-1), and sharp edges of dental enamel band two which are indicative of prevalence of tropical rainforest during this time interval (Appendix 2). Mean ordinated hypsodonty estimations in the present dissertation reveal that mean annual precipitation was 1500+ and mean annual temperature was >27C during early Miocene (Table 5.1). These findings are in accordance with the findings of Janis et al. (2004) that in environments where mean annual rainfall is > 1500 mm and high mean annual temperatures and more brachydont/browsing species are always found more (3-6) than the mixed-feeding ones (1-5), the type of vegetation is tropical rain forest. Early middle Miocene (Chinji Formation; 14.2-13 Ma, MN 4, 5) is marked by appearance of browsers (31-51%), forest frugivores (22.8-26.7%) including origin of Sivapithecus sp. and browse dominated mixed feeders (11.4-8.91%) (Tables 5.2, 5.3; Figures 5.5, 5.9). This ecomorphological composition reflects the closed ever green
vegetation showing abundance of C3 grasses with increasing proportions of savannas. Mid middle Miocene (13-12 Ma) is dominated by browsing taxa (50%) and forest
frugivores (26.8%) (Table 5.3; Figures 5.5, 5.9) which reflect closed C3 vegetation and
first appearance of C4 grasses at ca. 12 Ma portraying a dynamic shift from tropical evergreen forest to sub-tropical one. During late mid Miocene to early late Miocene (12- 11 Ma) shows decreasing proportion of browsing taxa (50-32.5%), closed habitat mixed feeders (18.9-13%) and increasing percentages of forest frugivores (26.8-35.1%) (Table 5.3; Figures 5.5, 5.9). This ecological continuum reveals the vegetational mosaics of sub- tropical forest, moist and dry deciduous forests with ecotonal habitats. During Middle Miocene, the mesowear signatures also exhibit the abundance of high and sharp dental cusps. However, the significant number of round and high cusps, in G. pinjabiensis, reveals the change in vegetation pattern from tropical evergreen forest to subtropical one
with the initiation of C4 grasses. The Chinji Formation (middle Miocene) of Lower Siwaliks documents the remains of fairly big leaves which represents a rich mesophytic 121
paleovegetation whereas the floral impressions recorded from Upper Siwaliks (Plio-
Pleistocene) are small and indicative of a dry or arid C4 grassland environment (Lakhanpal, 1970). In the ecosystem which recieve a mean annual precipitation range between 800 and 1400 mm, mean annual temperature range 20-25C, the percentage of browsing species between 50-32% whereas the proportion of mixed feeders range between 15-19%, the type of vegetation is humid evergreen forest (Figures 5.5, 5.9). These findings are in consonnance with the findings of Holdrige (1947, 1967), Janis et al. (2004), Eronen et al. (2010). Early Late Miocene (Nagri Formation 11-10 Ma) record comprises of decreasing percentage (32-29.5%) of browsers, forest frugivores (35.4-30.9%) and browsing mixed feeders (19-13.5%) and increasing proportion of grazing mixed feeders (2.70-11%) (Table 5.3; Figures 5.5, 5.9) which suggest closed vegetational regimes punctuating the essence of moist deciduous canopy forest with marked seasonality. During 10-9.00 Ma, presence of browsers (29.8-26%), forest frugivores (35.95%), browsing mixed feeders (13.51-11.9%) and grazing mixed feeders (7.7-13.6%) (Table 5.3; Figures 5.5, 5.9) led to the establishment of humidity-aridity gradient with humid and seasonal climate, closed
C3 vegetation and significant proportion of C4 grasses, moist deciduous forest, semi- closed temperate woodlands and initiation of dry deciduous forest (Table 5.4). In the habitat that recieve a mean annual precipitation range between 600 and 1000 mm, mean annual temperature range 18-22C, the percentage of brachydont browsing species ranging between (57-65%) while the number of closed habitat mixed feeders ranging between (23-28%) species reveals that the type of vegetation is moist deciduous forest (Tables 5.2, 5.3, 5.4). These findings are consistence with the findings of Janis et al. (2004) and Eronen et al. (2010). The 9-7 Ma interval shows decreaing of browsers (26-14.25%), forest frugivores (30.95-17%) and increasing of mixed feeders in open habitats (11.91-45.71%) and grazers (FG, GG) (2.35-5.65%) (Table 5.3). Mammalian fauna of early Late Miocene is comprised of forest frugivores, open habitat browsers, browse-dominated mixed feeders, and gaze-dominated mixed feeders; that infer the moist deciduous forest with widening of
C3 grasses and more increasing proportion of C4 grasses than the older intervals (Nelson, 2003). According to the present study the ecosystems where a mean annual precipitation 122
ranges between 600 and 800 mm, mean annual temperature ranges 15-18C, the number of brachydont browsers range between 14-26%, forest frugivores (17-31%), whereas the number of mixed feeders ranges between 28-48% and grazers (6.5%) (Tables, 5.2, 5.3, 5.4), the type of vegetation is temperate woodland. Other workers like Janis et al. (2004), Eronen et al. (2010) also support the above findings. Fossils of plants and pollen recovered from early to mid Late Miocene (18-8 Ma) deposits indicate the presence of moist tropical evergreen forests at lowlands, while the moist deciduous forests together with mosaics of pines at the higher altitudes (Hoorn et al., 2000; Osborne and Beerling, 2006). The floral progression during Late Miocene (8-6 Ma) revealed a significant increase in grasses with shrub pollen, and a declining of forest groups (Osborne and Beerling, 2006). All the previous studies or interpretations and the present investigation on ecomorphic data of Siwalik ungulates clearly reveals that,
despite of showing slight differences in the first emergence of C4 plants at temporal
resolution, the last 5 Ma displayed a dramatic C4 grasslands expansion in the Siwalik succession. According to Quade et al. (1995), this vegetational change appears to have permanent alterations in the dietary preferences of ungulates as well as ratids dwelling overbank/ near the rivers floodplains all across the Indian Siwaliks and most probably played a significant role in the major faunal turnover spanning from 8-7.5 Ma. The mid Late Miocene (8.5-6.5 Ma) hypsodont grazers appear to be coincided
with transition of C3 and C4 vegetation. It is also supported by Patnaik (2003). Data on this interval shows the existence of ecotonal grasslands, mosaics of woodlands, moist and dry deciduous forest portraying the picture of semi-closed vegetation. Change in preferred diets and habitats of taxa synchronized with the timings of origin and extinction events as taxa of closed habitats and frugivores went extinct first and open habitat taxa prevailed by later Late Miocene record. In the ecosystem which recieve a mean annual precipitation ranging between 500 and 700 mm, mean annual temperature range 11-15C the number of browsing species range between (33.5-16.5%), forest frugivores (14.3- 8.3%) whereas the number of mixed feeders range between (33.3-41.4%) species, grazers 7.2-16.6%, the type of vegetation is semi-humid savanna (Table 5.2, 5.3). Other workers like Holdrige (1947, 1967), Janis et al. (2004), Eronen et al. (2010), also ageee with these findings. 123
The 8-6 Ma interval shows a global increase in C4 plants as revealed by changes in the δ13C values of dental enamel of mammalian fossils in Asia (Siwaliks of Pakistan), South America, North America and Africa. This widespread and abrupt change can be linked to a decline in CO2 concentrations below the threshold that favoured C3- photosynthetic machinery in plants. The change initiated at lower latitudes because the
threshold value for C3 photosynthesizing plants was higher at warmer or higher temperatures (Cerling et al., 1997). It has been found that the species diversification and their range extension coincided with global vegetation changes in the latest Miocene (Bibi, 2007). Nevertheless, fossil plants are very rare from the Miocene deposits of the Potwar Siwaliks, stable isotopic values from paleosol carbonates provide evidence for vegetation composition on the flood plain (Badgley et al., 2008). The interpretation of floodplain paleosols carbon and oxygen isotope data from Siwalik of Pakistan indicates the presence of closed canopy forest before 7.5 Ma and increasing of open forest or woodlands and extensive grasslands afterwards (Quade et al., 1989) (Figures 5.3, 5.7). It is generally accepted that the Late Miocene paleovegetation changes in response to inception/ intensification of the South Asian monsoon system that in turn drive the faunal evolution in Siwalik succession. Mesowear signatures of an assemblage of 13 ungulate taxa from Late Miocene reveals that the average mesowear-III scores tend to be increased from 1.20-3.85 with junction points (JPs) from 1.33-3.78 throughout the Late Miocene (Appendix 2). Mesowear-II and mesowear-III analyses also reveal a gradual increase in mesowear scorings with high and sharp cusps becoming less frequent and increased prevalence of round, high, low and blunt cusps. These progressive shifts in mesowear signatures indicate a trend toward a decreased browse component and an increased graze component in the diet of Siwalik ungulates while retaining their mixed feeding strategies. This trend of increasing low and blunt cusps is consonance with the inference that forests and woodlands were replaced by open savannas and grasslands with warm season grasses as observed by Wolf et al. (2013) after their mesowear study of hipparionines from the Siwaliks of Pakistan. During Late Miocene, microwear pattern of two extinct suines and one tragulid show a high number of pits including large pits and predominantly mixed scratches. The 124
progressive occurrence of mixed scratches is indicative of gradient of C3 grasses and C4 grasses. Moreover, periodic emergence of fine scratches, coarse scratches and high number of pits suggest the progression of paleoseasonality that punctuated the decreased proportion of C3 grasses and increasing of C4 grasses throughout Siwalik succession. Microwear analysis of an assemblage of four boselaphines, an anthracothere, two rhinoceroses and a hipparionine horse shows the successively increasing number of scratches and decreasing number of pits which indicate the change in dietary shift from fruit to browse and from browse to graze. The changes in dietary and habitat preferences of these taxa reveal a change in vegetation from forest, woodland to savannas and grasslands during Siwalik succession. A large shift in carbon and oxygen isotope values during the late Miocene, initially documented in this Potwar Siwalik sequence, records changes in vegetation composition and structure linked to tectonic uplift of Himalayas and onset of the South Asian monsoon system (Quade and Cerling, 1995; Zhisheng et al., 2001; Behrensmeyer et al., 2007). This ecological assessment also illucidate the faunal changes during Late Miocene and Pliocene (Nelson, 2005). The carbon isotope signatures of paleosols from
Potwar Siwaliks of Pakistan revealed the first appearance of C4 grasses at about 7.7 Ma and thereafter complete dominance from 6.5 Ma onwards (Quade et al., 1989). Similarly,
the C4 grasss in Nepal Siwaliks first appeared at about 7 Ma (Quade et al., 1995) and in
some regions of the Indian Siwaliks indicated the first appearance of C4 grasses at ca. 6 Ma in the Subathu, Dehra Dun and Kangra sub-basins (Sanyal et al., 2004). The carbon isotope record demonstrates that after 8.1 Ma significant amounts of C4 grasses began to
appear and that by 6.8 Ma floodplain habitats included extensive C4 grasslands. Plant
communities with predominantly C3 plants were greatly diminished after 7.0 Ma, and
those with predominantly C4 plants, which would have been open woodlands or grassy woodlands, appeared as early as 7.4 Ma (Barry et al., 2002). Paleosol sequences of variable maturity indicate the presence of forest, woodland, and grassland vegetation that was stable for decades to thousands of years (Quade and Cerling, 1995; Behrensmeyer et
al., 2007). The transitional phase of vegetation or C3 to C4 vegetation was first explored in the Siwaliks of Pakistan by Quade et al., (1989) to be because of the rapid uplifting of Himalayas and orogenesis of Tibetan Plateau and later intensification of South Asian 125
monsoon system based on the advantages of comparison of C4 grasses with C3 plants under the monsoonal paleoclimates. However, Quade et al. (1995) further worked out that the link of monsoonal intensification with uplift of Himalayas and Tibetan Plateau is not straightforward and several cascades of uplifting events resulted in C4 grasslands expansion. Likewise, the Dorcatheriurn Y373 sp. (the larger tragulid taxon) is hypsodont and its succeeding taxon "Tragulidae/L101 sp." is even more hypsodont (Barry et al., 2002). This evidence is signifying a change from browse/fruigivorous in closed, semiclosed to mixed feeder or grazer in open one that support the change from C3 vegetation to C3-C4 transition followed by C4 grasslands. The 6-4.5 Ma interval chronicles proportion of browsers both (BB and HB) decreased from 14.25% to 10.24%, increasing proportion of MFO from 45.5% to 50% and that of grazers (FG and GG) 5.7-23% (Table 5.3) that suggest the essence of vegetation mosaics of deciduous forest, open woodlands and grassy savannas leading to birthing of modern monsoon system. In the ecosystem that recieve a mean annual precipitation 500-700 mm, mean annual temperature range 08-10C the number of brachydont browsers range between species (16.6-12%), forest frugivores (8.3-6%) whereas the number of open habitat mixed feeders range between (41.4-51.5%) species and grazers (16.6-19.4%, the type of vegetation is mosaics of grassy savannas and grasslands (Table 5.2, 5.3). These findings are in accordance with the findings of Holdrige (1947, 1967), Janis et al. (2004), Eronen et al. (2010). Fossil pollen records
indicate the complete disappearance of C3 wooded vegetation or forests at 5.5 Ma, and
consequent dominance of C4 grasslands on the Himalayan foothills or Siwaliks and the Ganges floodplain (Osborne and Beerling, 2006). The δ13C data compiled over the last
two decades document an expansion of C4 grasslands globally via the displacement of C3 vegetation with C3-C4 transition phase during the mid Late Miocene and followed by C4- vegetation during the Plio-Pleistocene (Cerling et al., 1997). During 4.5-2.5 Ma, the decreasing of browsers (10.24-7.04%), stable occurrence of MFO (48%) and increasing proportions of grazers (23-37%) (Table 5.3) shows more prevalence of grassy savannas and grasslands with declining mosaics of woodlands. It is
very much clear that the C4 grassland expansion was a global phenomenon initiating in middle Miocene and persisted to the recent ages. Soil carbonate analyses of the Pabbi 126
Hills sequence reveal that the vegetation was overwhelmingly C4 grasslands from 2.2 to 0.5 Ma (Quade et al., 1993). In the ecosystems that recieve a mean annual precipitation <500 mm, mean annual temperature range 06-8C, the proportion of browsing (HB, BB) species between 10-7%, range of forest frugivores is 8.3-6% whereas the percentage of open habitat mixed feeders range between 51.5-60% and that of grazers 25%, the type of vegetation is arid-adapted grassland (Table 5.2, 5.3). These findings are in consonance with the findings of Holdrige, (1947, 1967), Janis et al. (2004), Eronen et al. (2010). The substantial loss of vegetational diversity during Early Pleistocene coincides with the evidence drawn from analyses of paleosol carbonates that indicate dominance of C4 grasslands at this interval (Quade et al., 1993). The dominance of taxa >100 kg in the Upper Siwaliks is probably related to long-term climatic and vegetational changes in South Asia (Dennel et al., 2007). The vegetation models concerning Last Glacial Maximum (ca. 20 ka) indicate the overall spreading and dominance of grasslands utilizing C4 photosynthetic pathway because of decreasing pCO2 levels (Ehleringer et al., 1997; Cowling, 1999a, b). During 2.5-0.5 Ma, the decreasing of browsers (7.04-2.43%), increasing proportions of MFO (48-56%) and stable occurrence of grazers (35%) with disappearance of frugivores (Table 5.3) reveal the complete disappearance of forests/woodlands and dominence of grassy savannas and grasslands. It is an admitted premise that the mesowear, microwear and hypsodonty patterns among Siwalik ungulates had been changing in response to changes in the fibrous (phytolith and grit), and abrasive elements in the vegetation at which they have been feeding. Browsers take soft and less fibrous (little phytolith and grit-less abrasive) diet whereas mixed feeders and grazers incorporate more fibrous diets (grasses) which is subjected to more abrasive, tough and hard for their masticating machinery (Rensberger, 1973; McNaughton et al., 1985; Franz-Odendaal and Kaiser, 2003). Many workers are in agreement with the inference that a less fibrous diet leads to low teeth wear while a more fibrous food imparts higher dental wear as the greater volume of poor quality food must be ingested which necessitate more chewing (Damuth and Janis, 2011). The diverse habitat and dietary adaptations among Siwalik ungulates indicate diverse vegetational mosaics that deposit a wide variation in the occurrence and intensity of fibrous food, phytolith and the soil containing grit. The closed vegetation (forests, 127
woodlands) contains decreased amount of fibrous (phyolith and grit) food while semi- closed and open vegetation contains increased proportions of fibers, phytoliths and grit. Ingested soil (grit) is likely to be more abundant in the diet of ungulates than phytoliths and has a greater role in causing variation in dental wear. Such an interpretation is concordant with detailed relationships among hypsodonty, feeding spectrum, habitat and diet in extant ungulates and with the distribution of soil on plant surfaces and among habitats. The association between increased ungulate hypsodonty and a grazing food comes about because grazers are feeding on a particular grass (resource) in an open habitat and in a particular way i.e. near to the ground. The combination of these factors maximizes the amount of soil which the organism is likely to encounter and ingest unavoidably. Specialized grazers tend to ingest the highest levels of soil consumption, the highest values of hypsodonty, and the highest rates of dental wear among extant species contrary to mixed feeders and browsers (Damuth and Janis, 2011). The increasing magnitudes of fibrous food with phytolith, grit and soil consumption in ungulates diet punctuate changes in dental wear rates and increased hypsodonty that in return signal the changes in paleoclimatic and paleovegetational variables leading to exploration of paleoenvrinment of Pakistan Siwaliks. However, these abrasive elements in fibrous food do constrain the applications of mesowear, microwear and hypsodonty methods by increasing dental wear rates. To resolve this problem, slightly worn and unworn samples were selected and generated data were evaluated by comparing it with the standardized data of living analogues. Therefore, the variations in nature and amount of fibrous food, phytolith, grit, and geophagy did not have a significant effect on standards and observations of research protocols incorporated in this study as the unworn and slightly worn out samples (teeth) of ungulates were selected for mesowear, microwear and hypsodoty analyses. Moreover, for evaluation of results, the ecomorphic data of ungulates remains were compared with standardized data of their extant analogues.
128
5.2.1. DRIVERS FOR CHANGE FROM FOREST TO C3-C4 TRANSITION Impact of paleoclimate on faunal change or response of mammals to changing paleolimate has been signaling to explore diverse drivers that drove the forest to C3-C4 transition to C4 grassland dominance from ancient ages. Such drivers of environmental change can be explored and discussed from Siwaliks of Pakistan. Improved and comprehensive investigation of these diverse drivers and their mutual interactions has led
to frame the novel hypotheses concerning the initiations of C4 grasslands and has revived some of the old ones. All invoke local factors by focusing on increased aridity or shifts in ecological disturbance regime (Edwards et al., 2010). The 18O values, sedimentologic, and floral composition led to the establishment of seasonal paleoclimates with warm seasonal paleo-precipitation in the Siwaliks (Quade and Cerling, 1989; Zhisheng et al., 2001). The onset of dry and seasonal paleoclimate triggered the intensification of fire cycles in the Siwaliks which served as a driver of forest to C3-C4 transition or grassland transition and is favored by records of charcoal revealing increased incidents of fire in this region (Keeley and Rundel, 2003; Osborne and Beerling, 2006). The major problem associated with these scenarios of Late Miocene and Pliocene of Siwaliks is that they depend on mechanisms of modern ecosystems which maintain the
tree/shrub to grassland balance in C4-dominated ecosystems, however, the fossil record of Siwalik ungulates indicates that the great faunal transformation from forest to grassland
transition mainly documenting C3 grass dependent taxa (Figures 5.2, 5.3). Whether the
similar mechanisms drove the spreading of C3 grasslands is a contentious issue that remains to be resolved (Stromberg, 2005). Although, modern ecology elucidates that the abundance of woody vegetation versus grass is regulated by interplay of multiple factors at different spatio-temporal scales. Over the regional scale, grasslands are sustained by the interactions of climate, soil and ecological disturbance, punctuated by the traits of grasses and herbivorous fauna. Regionally, topographic factors generate variations in microhabitat which supports the mosaics of distinctive grass communities with diverse trait combinations (Coupland, 1992a, b, 1993). The general interpretation is that the traits which were, at best, indirectly
interrelated to C4 photosynthetic pathway, played important roles in permitting C4 grasses 129
to successively widen over vast areas/mosaics of natural grasslands during Late Miocene
of Pakistan that triggered the change from forest to C3-C4 transition during 8.5-6.5 Ma.
5.2.2. RATIONAL FOR DOMINANCE OF C4 GRASSLANDS
Studies on the most plausible causes that triggered the global expansion of C4 grasslands during the Late Miocene are still being actively debated. Various researchers have demonstrated that the Late Miocene expansion of C4 grasslands have long been coincided with the tectonically induced paleoclimatic changes, mainly brought about by
declining level of atmospheric CO2, large scale fire cycles, seasonality, intensification of
monsoon, and aridity. The factors that govern the progressive development of C4 grassland paleo-ecosystems in the Neogene involved increasing paleoseasonality at higher latitudes. This might have contributed to the great transformation of habitats from forest to grass dominated ones (Fox and Koch, 2004). Conversely, low seasonality is experienced in many present day low latitude areas mainly linked with the high temperatures (similar to those inferred from global pattern for species richness of mid Miocene) and different levels of rainfall. Therefore, it seems improbable that savannas of middle Miocene were existed outside the climatic spectrums that present day ecosystems constrain the presence of tropical savannas, and determine largely their associated potential productivity (Janis et al., 2004).
However, these major premises explaining Late Miocene C4 grasslands expansion seem improbable as the significant shortcomings linked with them, i.e.,
expansion/dominance of C4 grasslands in non monsoonal areas pose serious constraints on the hypothesis of intensification of monsoon. Similarly, aridity as being the common
driver of expansion of C4 grassland is not convincing because of non-substantial evidence that could suggest the onset of global arid conditions at ca. 9 Ma. The same geologic timescale and some most arid intervals such as the Triassic period, reflect no evidence of
C4 grasslands. The global pCO2 declining in the Late Miocene has been found to be the major driver (Singh et al., 2013). It is suggested that the initial declining of pCO2 <450 p.p.m.v created a paleoenvironment required for the initiation of C4 vegetation, however
the persistence of threshold value of pCO2 for a particular time in Late Miocene appears
to be the most probable cause of global C4 grassland expansion irrespective of knowing their time of first appearance. 130
The correlation of tectonic events such as uplift of Himalayas and Tibetan Plateau
with the levels of atmospheric pCO2 and vegetational changes reveals that it was actually the intense and continuous instability in the orogenesis of the Himalayas and Tibetan Plateau during Late Miocene may have trigerred the extensive chemical weathering of rocks of silica, which caused significant declining of atmospheric CO2 levels and that played a crucial role in changing the mode of photosynthetic processes and types of
paleovegetation. Late Miocene expansion of C4 grasslands in the Siwaliks was tectonically driven but was not by any global paleo-climatic changes. Conversely, Miocene uplifting of Himalayas might have been driving engine for global paleoclimatic
change that caused the expansion of C4 grasslands globally (Singh et al., 2013). However, this premise needs to be evaluated further by correlating the major tectonic events with global paleovegetational changes. Various workers have found that the existence of a strong monsoonal paleoclimate in the Siwaliks goes as back as ca. 16 Ma (Retallack, 1991) and ca. 10 Ma
(Kohn et al., 1999), therefore how can the absence of C4 grasses be elucidated during that
interval? The expansion of C4 grasslands in non-monsoonal areas of South and North
America also goes against the global expansion of Late Miocene C4 grasslands due to
monsoonal intensification. Physiological merits of C4 grasses over C3 grasses led Cerling
et al. (1997) to conclude that the atmospheric CO2 decline in low latitude temperatures in
Late Miocene was the main cause of C4 grasslands expansion. The pCO2 starvation or decline may have been a major factor for the initiation of C4 pathway; however, a time gap of ca. 10 Ma remains to be adjusted between major declines in the Oligocene CO2, and earliest current Late Miocene evidence of emergence of C4 grasses (Osborne, 2008). Singh et al. (2013) explored the monsoon intensifications at ca. 10, ca. 5, and ca. 1.8 Ma, however, the monsoonal intensification at ca. 10 Ma could not elaborate the
absence of C4 grasses at that interval. Moreover, some workers have considered the fire
to be an important driver in evolution and spreading of C4 grasslands during Late
Miocene (Scheiter et al., 2012). It is now admitted premise that the Late Miocene C4 grassland expansion was a major global phenomenon and lots of investigation in this regarding the most plausible driver for C4 grasslands expansion has been a contentious
issue. The efficiency of photosynthesis of C3 plants as compared to C4 grasses varies with 131
the levels of atmospheric pCO2 and growing seasonal temperature during daytime are
such that the C4 grasslands are flourished under low levels of pCO2 with elevated temperatures (Cerling et al., 1997; Morgan et al., 2011). Therefore, the first step to work out the probable cause for C4 grassland expansion would be to have either an elevation in temperature or a decline in atmospheric pCO2 values or the dominance of both from Miocene to Pleistocene/recent time (Singh et al., 2013).
The emergence of C4 photosynthetic pathway is not linked with a dramatic
decline in the level of pCO2 but could have probably been driven when pCO2 declined below the threshold value required for C3 pathway to occur (Singh et al., 2013). The
threshold value for concentration of pCO2 below which the C3 photosynthetic pathway could not be favored altogether, is turns out to be 450 p.p.m.v (Figure 5.3) that was
most probably achieved at ca. 9 Ma (Figure 5.2, 5.3). However, changes in pCO2 together with associated temperature do have relatioship with seasonality and precipitation patterns (humidity-aridity gradients) directly or indirectly. Keeping in view of fore mentioned premises, it is concluded that the Late Miocene intense and continuous instability in the orogenesis of the Himalayas and Tibetan Plateau caused significant declining of pCO2 which trigerred the change from C3-photosynthetic pathway to C4- photosynthetic one. The initial declining of threshold value of pCO2 <450 p.p.m.v in turn
created a paleoenvironment required for the initiation of C4 vegetation, however the
persistence of threshold value of pCO2 for a particular time in Late Miocene appears to
be the most probable cause of global C4 grassland expansion. The decline in pCO2 was accompanied by successive decrease in temperature and humidity which in turn reflected the increase in seasonality and aridity during Siwalik succession. The successive decreasing of pCO2 and temperature affected the faunal evolution directly whereas increasing seasonality and aridity impacted the faunal change indirectly. Combined interaction of these ecological drivers affected the immigration, speciation and extinction events of Siwalik mammals through times. Hence open habitat mammalian taxa (mixed feeders in open habitats and grazers) appeared and dominated at the expense of closed habitat taxa (browsers and mixed feeders in closed habitats) during late Miocene through Plio-Pleistocene of Pakistan Siwaliks. 132
Ecomorphic data collected during the present study reveals that C4 grasses first appeared around 12 Ma in Siwaliks of Pakistan and their proportion increased with the passage of time and expansion of C4 grasslands happened after 5 Ma due the persistence
of threshold value of (<450 p.p.m.v) pCO2 with its associated seasonality. All the previous studies or interpretations and the present investigation on ecomorphology of Siwalik ungulates clearly reveal that despite showing slight differences in the first
emergence of C4 plants at temporal resolution, the last 5 Ma displayed a dramatic C4 grasslands expansion in the Siwalik succession. According to Quade et al. (1995), this vegetational change appears to have permanent alterations in the dietary preferences of ungulates as well as ratids dwelling overbank/ near the rivers floodplains all across the Siwaliks and most probably it played a significant role in the major faunal turnover spanning from 8-7.5 Ma. 5.3. SPECIES DIVERSITY Mammalian species diversity varied considerably throughout the Siwalik series (18.3-0.5 Ma) in Pakistan. 5.3.1. SPECIES RICHNESS IN SIWALIK UNGULATES Early Miocene of Pakistan Siwaliks is represented by proboscideans, tragulids, bovids, suids, giraffids and rhinocerotids. All of these groups of mammals show very low and stable species richness (species diversity) in their respective communities till 16.5 Ma. (Table 5.2) Studies on artiodactyl (larger herbivorous mammals) and rodents (smaller herbivorous mammals) remains as representative of mammalian faunas, from 16-7 Ma on the resolution of 0.5 Ma reveals that the species richness was the highest in the middle Miocene (Chinji Formation), may be higher than that observed today, and then dramatically declined from 13.5 to 11 Ma onwards (Barry et al., 1990, 2002). Boselaphini: Species richness in boselaphines was low and stable (two species named Eotragus noyei, Paleohypsodontus zinensis) during Early Miocene and showed maximum diversity in Middle Miocene (Barry et al., 1991) and reached up to nine (9) species at 13.5 Ma and then gradually declined to 7 at 11 Ma and 6 at 8 Ma increased to 8 during 7- 6 Ma (Khan et al., 2009a). The number is reduced to half at 5-4 Ma followed by 3 during 3.5-2.5 Ma and again declined to 2 till 1.5 prior to their extinction (Table 5.2). 133
Bovini: Bovines first appeared in Siwaliks at 9.9 Ma (Late Miocene) with one representative. The number of species (richness) became two at 6.5 Ma that elevated to four at 3.5 Ma and attained maximum species richness (diversity) with eleven species during 2.5-1.5 Ma (Barry et al., 2002; Khan et al., 2009c, 2010b). Reduncini: The fresh grass grazers emerged at ca. 7 Ma with one species and three at 6.5 Ma and four at 5.5 Ma. By 3.5 Ma species richness increased from four to five followed by seven (7) during Pleistocene (Table 5.2) (Basu, 2004; Khan et al., 2008; Nanda, 2008). Antilopini: Antelopes first appeared in Middle Miocene and are represented by one species named Gazella sp. (Barry et al., 1991). The Late Miocene at ca. 10.5-8 Ma marks the existence of two species that became double in number from 8-6.5 Ma. These two antelopines appeared in the Siwalik record prior to the beginning of C3/C4 transition and might have had the narrow paleohabitat tolerances (Barry et al., 2002). The 6.5-4.5 Ma and 4.5-2.2 Ma intervals exhibit three and four species respectively (Basu, 2004; Nanda, 2008; Khan and Akhtar, 2014). Species richness of antilopini remains decreased from three to two in younger intervals (ca. 2.2-0.5 Ma) (Table 5.2). Hippotragini: One species of hippotragini appeared at ca. 6.5 Ma that persisted till 2.6 Ma and three species in the Pleistocene deposits have been documented (Table 5.2) (Basu, 2004; Nanda, 2008). Giraffids: The species diversity of giraffids was low and stable as Early Miocene record showed two (2) species of progiraffids, that were replaced by three species of true giraffids throughout Middle Miocene-Late Miocene-Early Pliocene (14-3.4 Ma) (Barry et al., 1991; Bhatti et al., 2012a, b) and two species named Giraffa sivalensis, Sivatherium giganteum were present during 3.5-0.5 Ma (Table 5.2) (Bhatti, 2005; Khan et al., 2011). Tragulids: Data shows that tragulids were represented by two species 18.3-17 Ma followed by three (3) during 16-12 Ma and increased to 5 around 11 Ma (Khan and Akhtar, 2013) and reduced to four (4) during 10-8 (Barry et al., 1991; Barry et al., 2002), thenceforth species decline continued from 3 to 2 whiles 8-7 Ma and 7-2 Ma intervals respectively (Table 5.2). Suids: The species richness is low and stable in Early Miocene suids as in the taxa of giraffids. The number of suid species increased to four at 14 Ma and six (6) at 13.5 134
(Samiullah et al., 2010) and then declined to four (4) and three (3) during 12-10 Ma and 9-2.5 Ma intervals respectively. The 2.5-1.5 Ma interval is represented by two species and is reduced to one whiles 1.5-0.5 Ma (Ahmad, 1995). Cervids: Five taxa of cervids appeared during ca. 5.3-4.5 Ma and declined to four (4) species at 1.5 Ma followed by three species at 0.5 Ma (Ghaffar et al., 2010, 2011). Rhinocerotidae: Aprotodon originated in Early Oligocene and diversified in Asia with Aprotodon fatehjangense evolving in Siwaliks until early Late Miocene (Beliajeva, 1954; Heissig, 1972; Wang, 1992). Aceratherium blanfordi was recorded from Middle Miocene (Chinji stratotype) of Siwaliks (Colbert, 1935; Heissig, 1972). In Middle Miocene, three more lineages named Brachypotherium, Chilotherium, Hispanotherium, Didermoceros were established. The temporal distribution of first two goes through Mid-Late Miocene of Siwaliks (Heissig, 1972). First two lineages belong to tribe Teleoceratini, Brachypotherium appearing close to the Diaceratherium of Europe and Chilotherium to the Aphelops of North America (Cerdeno, 1995). One species of Iranotheriine Hispanotherium is present in Middle Miocene (Cerdeno, 1996a; Inigo and Cerdeno, 1997) and Alicornopini, Aceratherini, Rhinocerotini are represented by one species each. Three (3) hypsodont species of Rhinocerotini are reported from 3.5 to 1.5/0.5 Ma (Table 5.2). Equidae: Cormohipparion sp. is a North American immigrant at 10.8 Ma that gave rise to Sivalhippus nagriensis at 10.3 Ma in the Siwaliks of Pakistan. Sivalhippus nagriensis was overwhelmingly abundant during 10.3-9.2 Ma with the exception of very rarely occurring Hipparion sp. small whiles 10-9.6 Ma interval. Sivalhippus theobaldi appeared at 9.4 Ma and had been rarely co-existing with Sivalhippus nagriensis for 0.2 Ma and then with the Sivalhippus perimense till 7.7 Ma. At 8 Ma Cremohipparion antelopinum and Sivalhippus anwari n. sp. emerged and initially co-existed with Sivalhippus perimense and were abundantly found for the period of 1.7 Ma (8-6 Ma) (Wolf et al., 2013). Sivalhippus anwari n. sp. and Cremohipparion antelopinum became less abundant in Early Pliocene and were replaced by two (2) species of Equus horses named Equus sivalensis and Equus sp. during the Upper Siwaliks (3.5-0.5 Ma) (Barry et al., 2002; Ghaffar, 2005; Dennel et al., 2006). 135
Anthracotheriidae: One species of anthracotheres appeared in early Miocene (18.3 Ma). Five species are documented from Middle Miocene deposits (13.9-11.2 Ma) that are declined to 4 during 11-9.3 Ma followed by three (3) in mid-late Miocene and one (1) in latest Miocene (Lihoreau et al., 2004, 2007). Proboscidea: One species of Gomphotherium lineage and two species of Deinotherium (Deinotheriinae) and one species of Synconolophus also originated at 16 Ma and persisted till 13 Ma. During Middle Miocene (14-11 Ma), Proboscideans show maximum species diversity and are represented by 10 species at 13.5 Ma that reduced to 6 at 11 Ma. Five species are recorded whiles 10-7 Ma followed by 4 during 6.5-5 Ma. Five (5) species of Proboscidea named Anancus sivalensis, Stegodon insignis, Stegodon sp., Elephus planiferons, Elephus hysudricus have been recorded during 3.5-0.5 Ma (Sarwar, 1974, 1977). Others: Chalicotherines and Doliochoerines are represented by one species each during 14-8 Ma interval as Hexaprotodon sivalense (Hippopotamidae) is, (6.10-2.20 Ma). Caprini is represented by one species during middle Miocene and one species (Sivacapra sivalense) during Late Plio-Pleistocene. One species of Alciphini; Demalops palaindicus is recorded from Pleistocene of Siwaliks (Barry et al., 2002; Badgley et al., 2005; Nanda, 2008). Species richness indices and species diversity are very less sensitive indicators, however, but still show interest (Barry et al., 2002). The number of species, as calculated by Fisher's α diversity index, shows a decline for large species of mammals by about 1/3 from the earlier values while small mammalian species α diversity declines by > 1/2 over the late Miocene. This decline represents a substantial reduction in the number of species, although it is not as drastic as indicated by the values of species richness themselves (Barry et al., 2002). The gradual decrease in species diversity of the mammalian fauna during the late middle Miocene of Siwaliks was real (Barry et al., 1990). Species diversity considerably varies showing a sudden rise in number of species between 15 and 13 Ma, followed by a decline in ruminant diversity after 12 Ma. Significant changes in relative abundance of taxa include an increase in bovids between 16.5 and 15 Ma, a decrease in tragulids after 9 Ma, and a very abrupt increase in murids at 12 Ma. Megacricetodontine rodents also decrease significantly at 12 Ma, and smaller declines 136
recorded among myocricetodontine and copemyine rodents after 16 Ma. An increase of dendromurine rodents at 15.5 Ma is also observed. Data on species richness and relative abundance (species diversity) of ungulates shows that the middle Miocene fauna of Pakistan Siwaliks displays large numbers of browsers (38) which exceed the number of browsers observed in today’s ecosystems and that is global phenomenon. However, after ca. 10 Ma, the numbers of browsing taxa gradually decreased to those exist in present day savannas. Conversely, the number of hypsodont taxa of ungulates increased gradually through the succeeding intervals of Siwaliks. Although they appear to dominate the Late Miocene temperate woodlands and savannas to more open habitat of Plio-Pleistocene yet they do not exceed their number experienced in present day habitats existing anywhere round the globe (Tables 5.2, 5.3). It is worth observing that some middle Miocene ungulate faunas of North America show large numbers of brachydont/browsers with 19 (a mean of 9.5) coexisting taxa (Janis et al., 2000, 2002) that are in excess of the numbers of brachydont/browsers observed in their extant analogous habitats (Janis et al., 2004). However, by ca. 10 Ma (Late Miocene), the number of browsing forms had declined to numbers consistent with those observed in today’s savannas. Hypsodont ungulates, in contrast, increase gradually in numbers through the Miocene, and although they dominate the ungulate forms of most of the Late Miocene and thereafter woodlands and more open ecosystems, yet they never increased the numbers of taxa seen in comparable extant habitats found anywhere in the world (Janis et al., 2004).
The factors that governed progressive development of C4 grassland paleo- ecosystems in the Neogene involved increasing paleoseasonality at higher latitudes. This might have contributed to the great transformation of habitats from forest to grass dominated ones. However, increasing seasonality can elaborate the decreases in species richness among ungulates throughout Miocene, but cannot explain why browsing taxa are superabundant in communities of Middle Miocene (Table 5.2, 5.3). Fox and Koch (2004) also agree with the above findings. Furthermore, low seasonality is experienced in many present day low latitude areas mainly linked with the high temperatures (similar to those inferred from global pattern for species richness of Middle Miocene) and different levels of rainfall. Therefore, it seems improbable that savannas of Middle Miocene 137
existed outside the climatic spectra that present day ecosystems constrain the presence of tropical savannas, and determine largely their associated potential productivity (Janis et al., 2004). 5.3.2. RODENT (MURID-CRICETID) SPECIES RICHNESS (DIVERSITY) THROUGH THE MIO-PLIOCENE IN THE SIWALIKS The murid-cricetid species diversity reveals that cricetids predominated during ca. 18.5 to 10.5 Ma. Around 10.5 Ma the cricetids and murids were represented by 8 species and 1 species respectively. During 9.5-8.5 Ma interval, both of these rodent groups were equal in number with three species each (Abudhabia, a gerbillid, appears at ca. 8.6 Ma) (Flynn and Jacobs, 1999). After 7.8 Ma murids began to dominate. Biochrnologic assessment reveals that Abudhabia extended its stratigraphic occurrence till the Pliocene spanning from 4 Ma to 2.5 Ma, where it gave rise probably to Tatera recorded at 2 Ma (Patnaik, 1997). By 6.3 Ma three (3) species of murid were recorded, of which, Mus, only being the first extant form. Murids diversified with documentary record of eight (8) species in Siwaliks at ca. 4 Ma. However, the greatest diversity happened at ca. 2.5 Ma (early Pleistocene) when about thirteen (13) species of murids emerged. The 2-1.8 Ma interval chronicles a slight decline of murid taxa to ten (10) in numbers (Figure 5.5). The reason for this decline may be taphonomic or sampling biases (Patnaik, 2003). These findings are in consonance with the findings made in the present dissertation. The findings of different workers regarding the mammalian biodiversity in Siwaliks of Pakistan are described as; Mammalian remains of Siwaliks chronicle community (diversity) stability during middle Miocene prior to 14 Ma, from 13.5 to 12.5 Ma, from 12 to 10 Ma, and from 9 to 8 Ma (Barry et al., 1990). The middle Miocene fauna is more stable, with species showing longer durations. The contrast in longevities corresponds with hypothesized greater environmental stability in the middle Miocene (Flynn et al., 1995). Most of the fossiliferous localities of Potwar Siwaliks document similar kinds of mammalian assemblages (Barry et al. 1982; 1990), whether temporally successive or geographically separated. This coherency in faunal association may reflect paleocommunity stability over substantial time intervals (Flynn et al., 1990a, b). Similarity of faunal composition over larger areas can be tested. The diversification in Middle Miocene happened during the interval of global cooling whereas the latest 138
Miocene decline in species diversity and increased faunal turnover accompanied by carbon and oxygen isotopic changes are coincided with globally increasing paleoseasonality and aridity (Barry et al., 1995). There is a trend of progressive increase in body size among bovids and giraffoids throughout the Siwalik sequence (Barry et al., 1990; Flynn et al., 1995). 5.3.3. FAUNAL TURNOVERS, RELATIVE ABUNDANCE AND CHANGE IN PALEOCOMMUNITY STRUCTURE Changes in mammalian species diversity through Siwalik series (18.3-0.5 Ma) explore the patterns of faunal turnover. Inferred first appearance datum and last appearance datum are the most sensitive indicators of faunal turnovers (Appendix 9). Ecomorphic data on Siwalik ungulates taxa showed the general evolutionary trends during Miocene with predominance of browsers, browsing mixed feeders and fruigivores that were replaced by mixed feeders in open habitats and grazers through Plio-Pleistocene (Table 5.2, 5.3). The mammalian faunal changes throughout the Miocene, from a browse-dominated fauna (brachydont ungulates) to mixed feeding/graze-dominated ones (mesodonts or hypsodont) has generally been interpreted as a simple faunal replacement, with the taxa of hypsodonts being better adaptive to the grasslands or open habitats (Shotwell, 1961). The general evolutionary trends during the Neogene are the successive replacement of more open habitat taxa, the slow demise of closed habitat adapted ones, and overall paleoclimate cooling (Janis, 1993). Early Miocene (18.3-14 Ma) mammalian fauna includes the appearance of forest frugivorous/browsing tragulid (Archoetragulus), two lineages of boselaphines (Eotragus noyie, Paleohypsodontus zinensis), two lineages of suids (Paleochoerus poscoei, Bunolistriodon sp.), 4 lineages of probocideans (Gomphotherium sp. Prodeinotherium sp., Deinotherium pentapotamie, Synconolophus ptychodus), 2 rhinocerotids (Brachypotherium fatehjangense, Brachypotherium perimense), and two giraffids (Progiraffa sp. Paleomeryx sp. 18.3-15 Ma) among browsers, two lineages of browsing rhinos such as Alicornops aff. laogouense (16.7-7.5 Ma) and Alicornops complanatum (16-12.5 Ma) with Chalicotherium salinum showed elevated turnover at 16 Ma. This ecological continuum is indicative of tropical and multicanopied rain forest accomodating arboreal community at upper canopies and terrestrial and overbank ones at ground level. 139
In the environments of dense canopy forest, the species richness of browsers did not seem to increase with gradually increasing levels of mean annual precipitation (MAP) and rainfall > 1500 mm/year, mean annual temperatures are high. Such ecosystem showed a tendency towards increase in canopy height and decrease in under-story vegetation with a corresponding decrease in available resources to browsing mammals. Therefore the number of browser species might be decreased as rainfall and mean annual temperatures are increased beyond such levels where tropical closed canopy forest supported the particular vegetation type (Janis et al., 2004). Middle Miocene (Chinji Formation) reflects important faunal turnover which suddenly elevates at 14 Ma reaches its peak at 13.5 Ma. This interval chronicles the appearance of two lineages of Sivapithecus, 9-10 browsing lineages of rhinos (i.e. Hispanotherium matritense, Gaindatherium browni, Aceratherium blanfordi, Chilotherium intermedium, Brachypotherium fatehjangense, Brachypotherium perimense etc.), 7-10 lineages of proboscideans (e.g., Anancus properimensis, Deinotherium indicum, Stegolophodon stegodontoides, Tetralophodon falconeri), 2 browsing suids (Listriodon pentapotamie, Conohyus sindiensis), 3 browsing giraffids (Giraffokeryx punjabiensis, Giraffa priscilla, Giraffa sp.), 3 frugivorous tragulids (Dorcatherium anthracotheroides, Dorcatherium “259 species” small, Dorcatherium sp. very small), 8 lineages of boselaphines depending on browse or mixed feeding in closed habitats (Sivaceros gradiens, Strepsipotax sp. Sivaceros eremita, Helicoportax praecox, Helicoportax tragelaphoides, Helicoportax sp. Miotragocerus gluten, Protragocerus gluten), a mixed feeding antelope (Gazella sp.) and 5 browse to fruit dependant anthracotheres (Anthracotherium punjabiensis, Merycopotamus dissimilis, Merycopotamus nanus, Hemimeryx sp., Microbunudon milaensis). The relative abundance of browsers (HB, BB; 54-50%), frugivores/selective browsers (23-29%), mixed feeders in closed habitats (11.4-12.5%) (Table 5.3; Figure 5.6) reveals that the prevalence of tropical evergreen forest and swampy habitats, with gilds of woodlands and savannas. These findings are in consonance with various paleoecologists. The pollen and wood fossil record in Nepal and Indian Siwaliks indicate the flourishing of tropical evergreen forest during Middle Miocene similar to the habitats of modern apes. These 140
forests were gradually replaced by moist deciduous forest followed by dry deciduous forest during late Miocene (Prasad, 1993; Corvinus and Rimal, 2001). The 13.5-11.5 Ma interval show the extinctions of a few browsing to mixed feeders in closed habitat (MFC) lineages i.e. Sanitherium sp., Alicornops laogouense, Helicoportax sp., appearance of forest dependant taxa; Gaindatherium browni, Sivaceros gradiens, together with persisting frugivorous, browsers and browse dominated mixed feeders belonging to proboscideans, anthracotheres and antelopes from earlier interval. The relative abundance of browsers (HB, BB; 50%-48.8%) decreased, slight increase in frugivores/selective browsers (29%-35%), increasing of mixed feeders in closed habitats from (12.5 % -18.9%), and first appearance of MFO (2.7%) (Table 5.3; Figure 5.6) reveal the existence of environmental mosaics of sub-tropical-moist deciduous forest with initiation of dry deciduous forest and savannas. The 11.5-10.5 Ma event indicates the extinction of selective browsers/fruigivores; Dorcabune anthracotheroides, the appearance of two new forms of frugivores/ browsers; Dorcabune nagrii, Dorcatherium “Y270 species”, Dorcatherium majus (Tragulids) (at 11.4-10.6 Ma), the persistence of 2 browsing lineages of giraffids (Giraffa priscilla, Giraffokeryx punjabiensis), and 10-8 rhinos. The occurrence and relative abundance of browsers (HB, BB) decreased from 48.8%-29.8%, whereas increasing of frugivores/selective browsers (25% -26%), mixed feeders in open habitats (MFO) (2.7%- 11.9%) and decreasing of mixed feeders in closed-habitats (18.9%-16.5%) (Table 5.3; Figure 5.6) suggest the existence of substantial mosaics of moist deciduous and increasing proportions of dry deciduous forest with patches of temperate woodlands and savannas. Turnover event at 10.3 Ma greatly differs from that of the two latest Miocene events with respect to the duration of the species involved. The disappearace of browsers; Giraffokeryx punjabiensis, Giraffa priscilla, Listriodon pentapotamie, and appearance of browsing to mixed feeding to grazing hipparionine horses (grazers), browsing and fruigivorous probocideans (Stegodon bombiferons Stegotetrabelodon sp., Synconolophus corrugatus, Tetralophodon sp.), mixed feeders in closed and open habitats; Tragoceridus sp., Miotragocerus gluten, Selenoportax vexillarius, Selenoportx lydekkeri (boselaphines), the fruigivores/selective browsers; Dorcatherium “311 sp”, Dorcatherium 141
“Y270” Dorcatherium majus (tragulids), rhinocerotids (Gaindatherium vidali, Chilotherium intermedium, occurrence of Brachypotherium fatehjangense, Brachypotherium perimense Chalicotherium salinum). The event at 10.3 Ma comprises taxa that were both common and of long duration, whereas the latest Miocene events include more taxa that were shorter ranging and less common. This is true of both the taxa that appear and those that disappear. That is, long enduring taxa both appear and go extinct at 10.3 Ma, whereas in the latest Miocene taxa of brief duration appear and disappear. The gradual decline of browsers (HB, BB) (29.8-26.2%), frugivores/selective browsers (35.4-31%), mixed feeders in closed habitats (17%-16%), increasing of MFO (11-11.9%), and first appearance of fresh grass grazers (2.38%) (Table 5.3; Figure 5.6) reveal the shrinking of moist and dry deciduous forest and widening of wooded savannas. According to Badgley and Behrensmeyer (1980), Late Miocene faunas from the Potwar Siwaliks showed 15 over 29 ungulates <100 kg. This implies that the existence of diverse niches of vegetation accommodating the significant number of browsers with abundant cover. A different mode of turnover develops between ca. 9-8 Ma interval that marks the extinction of 4 lineages of selective browsers (Schizochoerus, Microbunodon, Tatraconodon, Sivapithecus) and two fruigivorous lineages of tragulids, (Dorcabune nagrii, Dorcatherium “Y270”) and a browsing rhino (Gaindatherium vidali). This event displays the emergence of two grazing mixed feeders of boselaphines (Pachyportax sp. and Selenoportax sp.) 2 mixed feeding giraffes (Bramatherium taxa, Hydaspitherium sp.), two lineages of mixed feeding hipparionine horses (Sivalhippus theobaldi and Sivalhippus perimense), 2 MFC lineages of antelopes (cf. Prostrepsiceros vinayaki, Protragelaphus skouzesi), and one selelective browsing (SB) lineage of boselaphines (Miotragocerus salmontanus) (Table 5.2). The slight decrease of browsers (HB, BB), from 26.2%-23.3%, but significant decline of frugivores/selective browsers (31-16%), mixed feeders in closed habitats 17%-16.3%, and elevation of mixed feeders in open habitats (MFO) 11-27%, dry grass grazers; 4.64%-5.74% (Table 5.3; Figure 5.6) are suggestive of excessive fragmentation of moist deciduous forest and dry deciduous forest with increasing proportions of temperate woodlands-savannas. The forest fragmentation during this interval caused the extinction of Sivapithecus (Nelson, 2003, 2005). The 142
appearance of numerous very short ranging and rare species has also been reported (Barry et al., 2002). The extinct taxa are characterized by depleted values of oxygen and carbon isotopes that indicate their dependence on closed and humid forested habitat (Nelson, 2003). It is therefore a general feature of latest Miocene faunal change that contrasts to older periods, not just a peculiarity of the phase of more intense turnover. The data on body size together with species diversity and richness indices strongly suggest that the structure of mammal communities substantially changed after 10 Ma, and thenceforth at 7.8 Ma (Barry et al., 2002). The 7.8 Ma turnover is coincided with the change from equid dominated to even more balanced large mammalian assemblages. Emerging taxa are comprised of a high- crowned rodent and a hypsodont tragulid; Dorcatherium “373 sp.” (specimens of the taxon were recovered from fossil locality Y373 hence this name was assigned by Barry et
al., 2002), which incorporate C4 grasses in their diet and possibly reflect the existence of
C4 vegetation. The 7.8 Ma turnover events seem temporally coincided with the changes in depositional system of floodplain and vegetation (Barry et al., 2002). The turnover event at 7.8 Ma followed by 250 Kyr and the first paleosol carbon isotopic evidence
reveal that the widespread C4 vegetation on the floodplain might have concordant with other floodplain depositional changes (Behrensmeyer, 1987). The carbon isotopic evidence also shows that the habitats with C4 vegetation became very common on the Siwalik floodplain. This is very strong evidence for changing the availability of habitats and is synchronized with the emergence of mixed feeding or grazing taxa (Barry et al., 13 2002). In this event, the paleosol δ carbon signatures indicate that the habitats with C3 dominated plants still commonly persisted, thus elucidating the low rate of extinction (Badgley et al., 2008). The rhythmic momentum of decline of browsers (HB, BB) (23- 14.2%), frugivores/selective browsers (16%-17%), and mixed feeders in closed habitats (MFC; 16.3-2.85%), and successive progression of mixed feeders in open habitats (MFO; 27-45%), and grazers (FG, GG; 4.64-5.70%) (Table 5.3; Figure 5.6) is indicative of prevalence of woodlands with grassy savannas and limited mosaics of moist and dry deciduous forests. More extinctions occurred between 7.0 and 6.5 Ma including Propotamochoerus, an omnivorous suid which appear to have fed predominantly on fruit, nevertheless a few 143
individual specimens reflected the browsing diet; but the microwear analysis of two lineages of tragulids (Dorcatherium majus and Dorcabune nagrii) revealed the a diet comprising of browse as primary component and fruit being secondary, and Dorcadoxa, a mixed feeding bovid inhabiting in open woodlands with C4 component increasing through times (Nelson, 2003). Among the taxa analyzed in these studies, Hemimeryx and Tragoceridus do not until after 6.5 Ma. The relative abundance of browsers (HB, BB) (14.2-10.52%), frugivores/selective browsers (17.14-10.5%), mixed feeders in closed habitats (MFC; 2.85-2.63%), stable occurrence of MFO (37%-45%), and increasing proportions of grazers (FG, GG; 5.7-18.4%) indicate the existence of open woodlands and grassy savannas with limited patches of dry deciduous and moist deciduous forests (Table 5.3; Figures 5.2, 5.6). Turnover over during 6.5-5.5 Ma marks the appearance of fresh grass grazers (FG) “Reduncini D013 species”, (or Reduncini large sp.), a Hippopotamid (Hexaprotodon sivalense), and a Tragulidae 101 unnamed, the disappearance of an anthracothere (Hemimeryx sp.). The persistence of 2 lineages of proboscidea; (Stegodon bombiferons, Tetralophodon sp.), one anthracothere (Merycopotamus dissimilis), 2 lineages of rhinos (Chilotherium intermedium, Alicornops camplanatum), and Hipparionines, Dorcatherium nagrii, Giraffa punjabiensis, Tragoportax sp. Selenoportax sp., Miotragocerus large sp., Miotragocerus sp., Pachyportax latidens, Gazella lydekkeri, Antilope intermedia, Antilope planicornis. The relative abundance of browsers (HB, BB; 10.52-9.6%), frugivores/selective browsers (10.5-7.3%), disappearance of mixed feeders in closed habitats (MFC) and increasing proportions of MFO (44.7%-51.25%) and grazers (FG, GG; 18-21.7%) indicate the increasing proportions of open wooded and grassy savannas with limited mosaics of dry deciduous and moist deciduous forests (Table 5.3; Figures 5.2, 5.6). The 5.5-4.5 Ma interval shows the extinction of three browsing lineages (Eotragus, Miotragocerus gluten, Tragoportax sp.), one mixed feeding lineages of boselaphines (Selenoportax sp.), 3 browsing-mixed feeding lineages of rhinoceroses (Brachypotherium fatehjangense, Brachy. perimense, Chilotherium intermedium) and appearance of five taxa (two browsers and three MFO,s) of cervids, two FG bovines, a MFO boselaphine (Ruticeros pugio), three taxa of each of reduncini and antilopini, with 144
persistence of dry grass grazing taxa of Hipparion. The relative abundance of browsers (HB, BB; 9.6-10.24%), frugivores/selective browsers (7.6%), mixed feeders in open habitats (MFO; 51%), and grazers (FG, GG; 21.6-23%) reveals the prevalence of open- wooded and grassy savannas with rare patches of dry deciduous and moist deciduous forests. This composition is indicative of initiation of modern monsoon system. The faunal turnover during 4.5-3.5 Ma exhibits disappearance of hipprionine horses, Hydaspitherium-Bramatherium lineage, appearance of Antilope subtorta, Equus sivalensis, Equus sp. 3 taxa of hippotragini, 5 taxa of reduncini; persistence of cervids, Gazella lydekkeri, Antilope intermedia, Antilope planicornis (5.5-0.5), a bovine; Proamphibos lacrymans, boselaphines; 2-keel (a large-sized bovid), Pachyportax sp., Ruticeros pugio, Miotragocerus large sp., Hexaprotodon sivalense, Proboscidea; Stegodon bombiferons, Tetralophodon sp., an anthracothere; Merycopotamus dissimilis. The reconstruction of ecological composition based on presence and relative abundance of mammalian remains such as ever decreasing browsers (HB, BB;12%), frugivores/selective browsers (SB; 7.6%), and increasing of MFO (51%), and grazers (FG,GG; 23-28%) depict the prevalence of grassy savannas and grasslands with rare patches of dry deciduous and moist deciduous forests (Table 5.3; Figure 5.6). The significant faunal turnover between (mid Pliocene- Pleistocene 3.5-2.5) the Tatrot and Pinjor (Pabbi Hills) stratotypes reveals the reduction (3-2 taxa) in the taxa of Proboscidea, and more hypsodont grazer E. hysudricus (and S. cf. insignis) replaced E. planifrons, whereas Equus sivalensis, Equus sp. replaced Hipparion. There was appearance of D. palaeindicus and Hemibos triqueticornis (hypsodont bovids), Rhinoceros sivalensis, Rhinoceros sondaicus, Punjabitherium cf. platyrhinus, Sivatherium giganteum, and very common occurrence of hippotragini, reduncini, bovines and cervids. The reconstruction of habitat on the basis of presence of the mammalian taxa and their abundance revealed substantial changes whiles mid Pliocene to Pleistocene of Siwaliks in virtually disappearance of forest, widening of grassy savannas from about 64% to 70%, shrinking of wooded savannas ca. 32% - 19%, and expansion of desert from ca. 2% to 10% (Basu, 2004). Faunal turnover pattern during 2.5 to 0.5 Ma reflects the prevalence of mixed feeders in open habitats (MFO; 52-56%) and grazers (36.5%) with small proportions of 145
browsers and omnivores with complete extinction of fruigivores. Such ecological continuum is indicative of the predominance of grasslands and grassy savannas that replaced most of the forests (Table 5.3; Figure 5.6). Post 1.8 Ma events shows the disappearance of Sivatherium giganteum, Hexaprotodon sivalensis, anthracotheres, the large canid C. cautleyi and the hyaenid Hyaenictis or Lycyaena (Dennell et al., 2006) that marks a reduction in species diversity. Plio-Pleistocene habitats were most probably less diverse than that of the Miocene. Contributions of different workers concerning species diversity and community stability in Pakistani Siwaliks are chronicled as follows. A maxima in mammalian appearance was occurred at 13.5 Ma; and a high rate of appearances was sustained during 8.5 Ma to 7 Ma (mid Late Miocene) intervals. The significant peaks in last records were centered at 12.5 Ma, 9.5 Ma, and post-8 Ma. Assessment of pattern of species richness or size of mammalian assemblage suggests that the most impressive faunal turnover event resulting from coincident first and last appearance maxima began at 8 Ma and intensified at 7 Ma. This turnover episode includes both appearances and disappearances of taxa, but pace of extinction predominated at 9.5 Ma resulted the decline in species diversity. The peak of faunal appearances at 13.5 Ma was not accompanied by a high rate of disappearances; similarly the maxima of last records at 12.5 Ma were not accompanied by increased appearances. Both artiodactyls, particularly bovids, and small mammal (rodents) contribute to the early middle Miocene maxima and 9.5 Ma predominant extinction (with a gradual decrease in species richness in tragulids after 9 Ma and complete demise of its lineages around 2.2 Ma) during Late Miocene turnover (Barry et al., 1990, 2002). Late Miocene fauna is comprised of forest frugivores and browsers in semiclosed/open habitats throughout the record while most of the younger levels include mixed feeders and grazers in open habitats. Preferred diets and habitats of taxa correlate with the extinction, origination, and diversification at temporal resolution, as most of the forest frugivores and closed habitat species went extinct and then taxa of open habitat successively prevailed until the recent ages (Nelson, 2003). Fossil assemblages are considered as surrogates of paleocommunities. Any change in the repetition of successive mammalian fossil assemblage signifies the substantial change in paleocommunity (Flynn et al,. 1998). Fossil assemblage reveals that four types of paleocommunities such as 146
arboreal community, terrestrial macro-communities, terrestrial micro-communities with fossorial adaptations and stream and stream bank communities have been recognized in the Siwaliks. Vasishat (1978) also agrees with the above four assemblages. These communities had been varying in their frequency of occurrence at spatio-temporal scales (Figures 5.2, 5.6). CONCLUSIONS Ecomorphic data collected via hypsodonty (18.3-0.5 Ma), mesowear and microwear (13-5 Ma) approaches applied on 163 taxa of ungulate remains referred from 97 localities of Siwaliks of Pakistan offers powerful evidence for paleoclimatic and paleovegetational inferences and evolutionary interpretations. The cascade of tectonic events about the uplift of Himalayas and Tibetan Plateau may have been acting as driving engines for changes in climatic shift or ecomorphic adaptations of Siwalik ungulates. Mean ordinated hypsodonty and species composition reveal that the evolutionary progression of paleoclimate of Siwaliks changed from humid and warm (14-11 Ma) to seasonal climate with initiation of South Asian monsoon system during 11-9 Ma that intensified during mid-latest Miocene. That climatic regime was diferent from that of today’s patterns. Modern monsoon system emerged and replaced an earlier one at 5 Ma and intensified during the mid Plio-Pleistocene intervals. The multiple episodes of paleoclimatic changes triggered vegetation habitats to expand, shrink, or migrate across the landscape. The ecomorphic data and faunal composition also suggest that the vegetation configuration of Siwaliks changed from a closed-vegetation system
(prevalence of C3 vegetation) (18.3-8.5 Ma) to a semi-closed one and from that (C3-C4 transitional vegetation) (8.5-6.5 Ma) to to an open vegetation system (abundance of C4 vegetation) (6.5-0.5 Ma). The persistence of threshold value of pCO2 for a particular time in Late Miocene appears to be the most probable cause of global C4 grassland expansion.
The decline in pCO2 was accompanied by successive decrease in temperature and humidity which in turn reflected the increase in seasonality and aridity during Siwalik succession. The herbivorous species dependant on particular vegetation or trophic resources shifted their biogeographic ranges to occupy the preferred habitats that in turn stimulated speciation, immigration, extinction events within mammalian paleocommunities though times. Integrated analyses of these processes led to the 147
emergence of long term patterns of faunal turnover, species diversity and paleocomminty structure in the Siwaliks of Pakistan. Change in paleocommunity structure of Siwalik ungulates, rodents and hominoids indicate that species diversity and increased episodes of faunal turnovers in Late Miocene are concordant with global climatic regimes of increased aridity and paleoseasonality. Species richness and relative abundance/species diversity of the browsing ungulate remains was highest (38 in number) the middle Miocene of Pakistan Siwaliks which exceed the number of browsers observed in today’s ecosystems and that was global phenomenon. After ca. 10 Ma, the numbers of browsing taxa gradually decreased. Conversely, the number of hypsodont taxa of ungulates increased gradually through the succeeding intervals of Siwaliks. Although they appear to dominate the Late Miocene temperate woodlands and savannas to more open habitat of Plio-Pleistocene yet they did not exceed in number the present day habitats. The 18.3 to 15 Ma intervals present the dominance of forest frugivores/selective browsers and browsers suggest the existence of dense multicanopied rain forest. In the Middle Miocene, the sudden increase in species richness with relative abundance of browsers, frugivores/selective browsers and mixed feeders in closed habitats reveals that the prevalence of tropical evergreen forest and swampy habitats at 13.5 Ma. Mid-Middle Miocene (13-12 Ma) reflect the predominance
of browsers and forest frugivores that indicate closed C3 vegetation and first appearance
of C4 grasses depicting a change in vegetational shift from tropical evergreen forest to sub-tropical one. The 12-11 Ma interval chronicles decreasing proportion of browsers, increasing percentages of frugivores and browsing mixed feeders that reveal the mosaics of sub-tropical forests, moist and dry deciduous forests with ecotonal habitats. The 9-8
Ma interval represents increasing mosaics of C3-C4 transitions reflect slight decrease of browsers, but significant decline of frugivores/selective browsers, mixed feeders in closed habitats, and elevation of mixed feeders in open habitats, and dry grass grazers are suggestive of excessive fragmentation of moist deciduous forest that caused extinction of Sivapithecus and that of dry deciduous forest with increasing mosaics of temperate woodlands and savannas. During 8-6.5 Ma interval, semi-closed vegetation system provided platform for great faunal transformation from closed habitat browsing and mixed feeding to that of 148
open ones with progressive influx of grazers. Biostratigratigraphic data reveal that most
of species reliant on C3 vegetation disappeared during the C3-C4 transition to C4 vegetation whereas immigrants were grazing mixed feeders or grazers. The frugivores and browsers, who changed their diets by incorporating graze, persisted and then switched to C4 grasses. The significant number of long lasting lineages belonging to group hipparionine horses, rhinoceroses, boselaphines, sivatherines, antelopes and a tragulid altered their feeding preferences from browsers to mixed feeders in open habitats/grazers. However, more sampling and investigation is required to support this premise. During 6.5-4.5 Ma interval, the further decrease of browsers and frugivores and successive increase of mixed feeders, and grazers reveals the prevalence of open wooded and grassy savannas with rare patches of dry deciduous and moist deciduous forests. Faunal turnover during 4.5-3.5 Ma exhibits the ever-decreasing browsers, frugivores/selective browsers, and increasing of open habitat mixed feeders, dry grass grazers that depict the prevalence of grassy savannas and grasslands with rarest mosaics of dry and moist deciduous forests. Turnover episodes in 2.5-0.5 Ma reflect the prevalence of mixed feeders in open habitats and grazers, the small proportions of browsers and omnivores with complete extinction of fruigivores. Such ecological continuum suggests the predominance of grasslands that replaced most of the forests. On the basis of mean hypsodonty estimations, this study presents more precise estimation of paleoclimatic variables such as precipitation patterns and temperature ranges than the earlier ones. However, further compilation of data from other fossil localities of this region may reveal even more precise precipitation estimation than the present study. A new method of mesowear (Type III) introduced in this study reveals better dietary inferences of Siwalik ungulates than the previous ones. It is a great contribution to the understanding of mammalian paleoecology. Hence this study provides a comprehensive account of paleoenvironment of Pakistan Siwaliks in relation to mammalian biostratigraphic and paleoecologic processes at an evolutionary scale.
149
Table 5.1; Infrences of mean annual precipitation from mean ordinated hypsodonty values (listed in Table 4.17) by correlating with extant communities.
Age MOH MAP M A T Paleoclimate (Ma) (mm) ranges
18-15 <1 1500+ > 27C Planetary (warm &humid) 15-12.5 1.0-1.2 1200-1500 22-26C Warm, Humid and uniform (incipient aridity) 12.5-11 1.2-1.4 800-1200 20-24C Subtropical, 11-9 1.4-1.8 600-1000 18-20C Seasonal, initiation of South Asian monsoon system 9-8 1.8-2.2 700-800 16-18C Pronounced seasonality of rainfall 8-5 2.2-2.6 500-700 11-15C Intensification of Asian monsoon system 4-3 2.6-2.8 300-500 8-10C Development of modern monsoon system
3-0.5 2.8-3.0 200-300 06-08C Intensification of modern monsoon system Key to symbols; MOH= mean ordinated hypsodonty, MAP= Mean annual precipitation (mm), MAT= mean annual temperature (following Janis et al., 2004; Eronen et al., 2010) 150
Table 5. 2; Species richness in ungulate remains recorded from Siwaliks of Pakistan. Age (Ma) Age (Ma) Proboscideans Anthratheres Hippotragini Boselaphini Hominoids Antilopini Reduncini Tragulids Giraffids Cervids Rhinos Equids Bovini Suidae Others Others Total
0.50 0 02 02 02 - 02 - 02 - 02 06 05 01 02 03 29 1.00 0 02 03 02 - 02 - 02 - 02 06 05 01 02 03 29 1.50 0 03 03 02 - 02 - 05 02 02 06 05 02 02 03 35 2.00 0 03 03 02 - 02 - 05 03 02 09 05 02 02 03 39 2.50 0 02 03 02 02 02 - 05 03 03 09 06 03 01 03 43 3.00 0 02 03 02 02 - - 05 03 03 07 05 03 01 01 36 3.50 0 02 02 02 02 - 01 05 03 03 07 05 03 01 01 37 4.00 0 03 02 02 02 03 01 05 04 03 02 03 03 01 01 35 4.50 0 04 02 02 02 03 01 05 04 03 02 03 03 01 01 36 5.00 0 06 02 02 02 03 01 05 04 02 02 03 03 01 01 37 6.00 0 06 03 02 02 04 01 - 08 02 02 02 03 01 01 37 6.50 0 06 03 02 03 04 02 - 08 02 02 02 03 - 01 38 7.00 0 08 04 02 05 04 02 - 08 02 01 - 03 - - 39 8.00 0 08 05 03 05 04 02 - 06 03 01 - 03 01 - 41 9.00 2 07 05 02 08 03 03 - 07 02 01 - 03 01 - 44 10.0 2 07 05 02 08 03 04 - 07 02 - 03 01 - 44 151
Table 5.2 (continued) Age (Ma) Age (Ma) Anthracotheres Anthracotheres Prboscideans Hippotragini Boselaphini Antelopini Reduncini Hominids Tragulids Giraffids Cervidae Rhinos Equids Bovini Suidae Others Others Total
11.0 1 07 08 - 09 02 04 - 08 01 - - 05 - - 45
11.5 1 08 10 - 05 02 05 - 07 01 - - 06 01 - 47 12.0 1 08 10 - 06 03 05 - 07 01 - - 06 01 - 48 12.5 1 08 10 - 06 03 05 - 09 01 - - 06 01 - 50 13.0 1 09 10 - 06 03 05 - 09 01 - - 06 01 - 51 13.5 1 10 10 - 06 05 05 - 09 01 - - 06 01 - 54 14.0 0 07 09 - 06 04 02 - 08 01 - - 06 01 - 45 15.0 0 04 05 - 02 02 02 - 02 - - - 02 - - 19 15.5 0 04 04 - 02 02 02 - 02 - - - 02 - - 18 16.0 0 04 04 - 02 02 02 - 02 - - - 02 - - 18 16.5 0 04 04 - 02 02 01 - 02 - - - 02 - - 17 17.0 0 04 03 - 02 02 01 - 02 - - - 02 - - 16 17.5 0 03 03 - 02 02 01 - 02 - - - 02 - - 15 18.0 0 03 02 - 02 01 01 - 02 - - - 02 - - 13 18.3 0 03 02 - - - 01 - 02 - - - 02 - - 10 Others=Hippopotamidae, Alcephini, Caprini.
152
Table 5.3; Relative abundance among different ungulate groups from Pakistan Siwaliks % Fr-SB %MFO %MFO %MFC %MFC Int. Ma % HB % HB Fr/SB %GG %GG MFO %BB MFC %FG %O %O GG GG HB HB BB BB FG O
0.50 1 - - 2 - 23 6 09 2.43 0 0 4.86 0 56.09 14.63 21.9
1.00 1 - 1 2 - 24 6 10 2.27 0 2.27 4.54 0 54.54 13.63 22.7
1.50 2 - 1 2 - 26 6 10 4.25 0 2.12 4.25 0 55.13 12.76 21.5
2.00 2 - 1 2 - 25 7 11 4.16 0 2.08 4.16 0 52.08 14.58 22.9
2.50 2 2 2 2 - 24 8 11 3.52 3.52 3.52 3.52 0 47.05 15.68 21.6
3.00 2 2 2 4 - 19 6 5 5.0 5.0 5.0 10.0 0 47.5 15.0 12.5
3.50 2 2 2 4 - 18 6 5 6.89 6.89 6.89 10.25 0 46.15 15.38 12.8
4.00 2 3 2 4 - 19 6 3 5.12 7.69 5.12 10.25 0 48.71 15.38 7.69
4.50 2 3 2 4 - 19 6 3 5.12 7.69 5.12 10.25 0 48.71 15.38 7.69
5.00 2 3 2 4 - 21 6 3 4.87 7.32 4.87 9.75 0 51.21 14.36 7.32
6.00 2 4 2 5 1 17 5 2 5.26 10.52 5.26 13.16 2.63 44.74 13.16 5.26
7.00 2 6 3 5 1 16 1 1 5.71 17.14 8.57 14.28 2.85 45.71 2.85 2.85
8.00 7 7 3 5 7 12 1 1 16.3 16.3 7 11.63 16.3 27.90 2.32 2.32
9.00 9 13 2 5 7 5 1 - 21.43 30.95 4.8 11.90 16.7 11.90 2.38 - 153
Table 5.3 continued…… BB % Fr-SB %MFO Int. Ma %MFC % HB Fr/SB %GG MFO %BB %FG MFC %O GG HB HB FG O
10.0 9 13 2 4 5 4 - - 24.32 35.14 5.5 11 13.51 11 0 0
11.0 10 13 2 4 7 1 - - 27.02 35.1 5.5 11 18.9 2.70 0 0
12.0 17 11 3 4 6 - - - 41.5 26.83 7.31 9.75 14.6 0 0 0
12.5 17 11 4 4 6 - - - 40.5 26.2 9.5 9.5 14.3 0 0 0
13.0 19 12 4 4 6 - - - 42.2 26.7 8.9 8.9 13.4 0 0 0
13.5 19 14 5 4 6 - - - 39.6 29.16 10.42 8.33 12.5 0 0 0
14.0 14 8 5 4 4 - - - 40 22.8 14.3 11.4 11.4 0 0 0
15.0 5 3 4 3 3 - - - 27.8 16.7 22.2 16.7 16.7 0 0 0
16.0 7 3 4 2 3 - - - 36.8 15.8 21.05 10.53 15.8 0 0 0
17.0 5 3 3 2 1 - - - 35.7 21.4 21.4 14.3 7.14 0 0 0
18.0 5 2 3 2 1 - - - 38.5 15.4 23.07 15.4 7.7 0 0 0
18.3 3 2 2 2 1 - - - 30 20 20 20 10 0 0 0
154
Table 5.4; Summary of environmental evolution of Pakistan Siwaliks in relation to changes in paleoclimate, vegetation, and paleocommunity structure
155
FIGURES
Figure 5.1; Neogene record of CO2 and temperature change, including evidence for Arctic glaciation, fossil and molecular dating evidence for grass evolution, and charcoal records. Plt., Pleistocene; Plio., Pliocene; K, Cretaceous; C, Chloridoideae; Pan, Panicoideae; Ar, Arundinoideae; An, Andropogoneae; B, Bambusoideae; P, Pooideae; BEP, Bambusoideae-Ehrhartoideae-Pooideae; PAC, PACMAD (taken and modified from Edwards et al., 2010).
Figure 5.2; Neogene record of changes in vegetation structure (from forest vegetation to an environment with substantial amount of grass cover), and of photosynthetic pathway (C3 versus C4), in vegetation and in diet of ungulates from various regions. Ol= Oligocene, Plt=Pleistocene (taken and modified from Edwards et al., 2010). 156
Figure 5.3; Predicting C3/C4 dominance of grasses related to temperature and partial pressure of
CO2 according to which photosynthetic pathway has the greater quantum yield; here ‘temperature’ is the daytime growing-season temperature (adapted from Cerling et al., 1997).
Diet and Habitat Type Figure 5.4; Relationship of mean hypsodonty index (HI) to diet and habitat type based upon 133 species of living ungulates of known dietary and habitat preference. Grazer = ≥ 90% grass in the diet; Mixed/Grazer (Mixed/G) = 50–89% grass in the diet; Mixed/Browser (Mixed/B) = 11–49% grass in the diet; Browser = ≤ 10% grass in the diet. Forest = closed forest, with few or no clearings; Woodland = more open forest where canopy is still mostly continuous but ground cover may include grass; Savanna = open habitat with grass and scattered trees and bushes; Grassland = grassland steppe with no significant woody plant cover. Height of bars indicates mean HI, and error bars show ±1 standard error (Source; Damuth and Janis, 2011). Succcession of living communities has been taken as model for characterization of mammalian fossil communities during Siwaliks series as mentioned in Figure 5.5. 157
5 Grasslands Savannas Woodlands Forests
4 BR MFC 3 MFO Gr
2
Mean Hypsodonty Index Hypsodonty Mean 1
0 0.5 1 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 18.3 Age (Ma) Diet and Habitat Preferences Figure 5.5; Relationship between mean hypsodonty and evolution of paleocommunities and paleovegetation in the Siwaliks of Pakistan
Figure 5.6; Carbon isotope record from pedogenic carbonates (Quade and Cerling, 1995; Behrensmeyer et al., 2007) and mammalian dental enamel (Morgan et al., 1994; Nelson, 2007) show a pronounced shift to lighter values starting 8.5 Ma. Over the well sampled interval from 9.5 to 7.0 Ma, mammalian 13C values vary more than do paleosol values, indicating that mammalian herbivores sampled a broader range of habitats than documented by the paleosol record. For ease of comparison of paleosol and mammal 13C values with respect to vegetation, all paleosol values were adjusted by -1‰ because the fractionation values (vegetation to palesol carbonate, vegetation to mammalian enamel) differ by this amount (source; Badgley et al., 2008). 158
Figure 5.7; Species richness (diversity) in ungulate remains through the Siwaliks series of Pakistan.
Figure 5.8; Number of murid and cricetid rodent species found or inferred to be present between 18 and 1.5 Ma in the Siwalik sediments. Data representing 18 to 5.7 Ma of Pakistan (Flynn et al., 1995), 6.5 Ma of India ( Flynn et al., 1990a, b; Tiwari, 1996), 4 Ma of India (Patnaik, 1997), 2.5 Ma of India (Patnaik, 1997; Gupta and Prasad, 2001), 2 Ma of India (Raghavan, 1990; Patnaik, 1997), 1.8 Ma of India (Patnaik, 1997), and of Pakistan (Jacobs, 1978; Musser, 1987) (Source: Patnaik, 2003). 159
70 %B % Fr 60 %MFC %MFO 50 %GR
40
30
20 Percentage of ecomorphic characters ecomorphic of Percentage
10
0
Age (Ma)
Figure 5.9; Relative abundance among ungulates and succession of paleocommunities during Siwalik chronology. Abbreviations; B=browsers, Fr= frugivores, MFC= mixed feeders in closed habitat, MFO= mixed feeders in open habitat, GR= grazers.
160
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Appendix 1; Locality database of mammalian fossils from Siwaliks of Pakistan
Sr# Locality Mid point Upper Lower Age No of Assigned age age spread intervals Interval 1 SRDK-73 ca.1.2 - - - - ca.1.2 2 SRDK-642 ca. 1.4 - - - - ca. 1.4 3 SRDK-362 ca. 1.9-1.7 - - - - ca. 1.9-1.7 4 KhS ca.2.0 - - - - ca.2.0 5 PirG ca. 2.0 - - - - ca.2.0 6 JK ca.2.5 - - - - ca.2.5 7 B0139 3.30 3.005 3.596 0.591 07 3.30 8 B122 3.50 - - - - 3.50 9 Kakrala 3.50 - - - - 3.50 10 KTLKND 3.50 - - - - 3.50 11 Dndibatian 3.50 - - - - 3.50 12 RTS 3.50 - - - - 3.50 13 DKAWN 3.50 - - - - 3.50 14 B044 4.00 - - - - 4.00 15 B119 4.00 - - - - 4.00 16 B147 4.50 - - - - 4.50 17 B120 4.50 - - - - 4.50 18 B150 5.50 - - - - 5.50 19 B151 5.50 - - - - 5.50 20 B152 5.50 - - - - 5.50 21 B157 5.50 - - - - 5.50 22 RTL-1 ca. 6.00 - - - - ca. 6.00 23 B109 6.196 6.122 6.269 0.147 03 6.20 24 B118 6.385 6.290 6.480 0.190 03 6.40 25 B110 6.392 6.303 6.480 0.177 03 6.40 26 B112 6.392 6.303 6.480 0.177 03 6.40 27 B008 6.488 6.306 6.670 0.364 05 6.50 190
Appendix 1 (continued) Sr# Locality Mid Upper Lower Age No of Assigned point age age spread intervals Interval 28 B024 6.488 6.306 6.670 0.364 05 6.50 29 B135 6.488 6.306 6.670 0.364 05 6.50 30 B006 6.566 6.448 6.684 0.236 04 6.60 31 B007 6.566 6.448 6.684 0.236 04 6.60 32 RTL-2 ca. 7.00 - - - - ca. 7.00 33 B117 7.222 7.083 7.360 0.277 04 7.20 34 B104 7.863 7.802 7.924 0.122 02 7.90 35 B114 7.960 7.592 8.327 0.735 08 8.00 36 B043 7.973 7.896 8.049 0.153 02 8.00 37 B046 8.005 7.961 8.049 0.088 01 8.00 38 B025 8.088 7.802 8.374 0.572 07 8.10 39 B026 8.088 7.802 8.374 0.572 07 8.10 40 B027 8.088 7.802 8.374 0.572 07 8.10 41 B028 8.088 7.802 8.374 0.572 07 8.10 42 B029 8.088 7.802 8.374 0.572 07 8.10 43 B030 8.088 7.802 8.374 0.572 07 8.10 44 B031 8.088 7.802 8.374 0.572 07 8.10 45 B032 8.088 7.802 8.374 0.572 07 8.10 46 B033 8.088 7.802 8.374 0.572 07 8.10 47 B034 8.088 7.802 8.374 0.572 07 8.10 48 B035 8.088 7.802 8.374 0.572 07 8.10 49 B036 8.088 7.802 8.374 0.572 07 8.10 50 B037 8.088 7.802 8.374 0.572 07 8.10 51 B103 8.125 8.012 8.238 0.226 03 8.10 52 B106 8.125 8.012 8.238 0.226 03 8.10 53 B040 8.151 8.065 8.237 0.172 02 8.20 54 B038 8.270 8.226 8.313 0.087 02 8.30 191
Appendix 1 (continued) Sr# Locality Mid Upper Lower Age No of Assigned point age age spread intervals Interval 55 B044 8.270 8.226 8.313 0.087 02 8.30 56 B105 8.270 8.226 8.313 0.087 02 8.30 57 B042 8.300 8.226 8.374 0.148 03 8.30 58 B041 8.331 8.287 8.374 0.087 02 8.30 59 B045 8.331 8.287 8.374 0.087 02 8.30 60 B111 8.760 8.247 9.272 1.025 12 8.80 61 B115 8.760 8.247 9.272 1.025 12 8.80 62 RTL-3 ca. 9.00 - - - - ca.9.00 63 B009 ca. 10.0 - - - - ca. 10.0 64 B010 ca. 10.0 - - - - ca. 10.0 65 B011 ca. 10.0 - - - - ca. 10.0 66 B012 ca. 10.0 - - - - ca. 10.0 67 B013 ca. 10.0 - - - - ca. 10.0 68 B017 ca. 10.0 - - - - ca. 10.0 69 B018 ca. 10.0 - - - - ca. 10.0 70 B019 ca. 10.0 - - - - ca. 10.0 71 B020 ca. 10.0 - - - - ca. 10.0 72 B021 ca. 10.0 - - - - ca. 10.0 73 B022 ca. 10.0 - - - - ca. 10.0 74 B023 ca. 10.0 - - - - ca. 10.0 75 B107 ca. 10.0 - - - - ca. 10.0 76 B108 ca. 11.0 - - - - ca. 11.0
77 B137 ca. 11.0 - - - - ca. 11.0
78 B155 ca. 11.0 - - - - ca. 11.0
79 B159 ca. 11.0 - - - - ca. 11.0
80 B0123 11.85 11.955 11.750 0.205 - 11.80
192
Appendix 1 (continued)
Sr# Locality Mid Upper Lower Age No of Assigned point age age spread intervals Interval 81 B0054 12.235 12.108 12.355 0.247 - 12.25 82 B0057 12.664 12.543 12.785 0.242 - 12.70 83 B0055 12.664 12.543 12.785 0.242 - 12.70 84 B0056 13.04 12.893 13.200 0.493 - 13.00 85 B0125 13.355 13.223 13.487 0.264 - 13.40 86 B0051 13.751 13.672 13.830 0.158 - 13.75 87 B0048 13.808 13.760 13.857 0.097 - 13.81 88 B0049 13.808 13.760 13.857 0.097 - 13.81 89 B0050 13.808 13.760 13.857 0.097 - 13.81 90 B0126 13.824 13.705 13.944 0.239 - 13.83 91 B0047 13.842 13.783 13.901 0.218 - 13.85 92 B0127 14.036 13.983 14.095 0.112 - 14.00 93 B0052 14.01 13.923 14.106 0.183 - 14.00 94 B0058 14.208 14.180 14.236 0.056 - 14.21 95 B0053 14.246 14.210 14.282 0.072 - 14.25 96 Jaba - - - - - ca. 15 97 DML - - - - - ca. 16
Key to Symbols: B=Branum Brown, SRDK=Surdhok, JK=Jarikus, RTS=Rohtas, DKAWN=Dhokawan, KTLKND=Kotalkund, PirG=Pir Ghaffar, KhS= Khurla sharif, DML=Domeli.
193
Appendix 2; Mesowear Type III data of living and fossil species of mammals.
Species Days M-II M-III AB P B J P First goat experiment Capra hircus browsing 20 1 1.5 1.5 2 2.5 Capra hircus browsing 19 2 1.35 1 1.75 2 Capra hircus browsing 19 2 1.5 1.5 1.5 2.75 Capra hircus browsing 18 2 2 1 3 2.5 Capra hircus browsing 19 2 2.25 2 2.5 3 Average 1.8 1.72 1.3 2.15 2.50 Capra hircus grazing 20 2.25 3 3 3 3 Capra hircus grazing 19 2 3 3 3 2.5 Capra hircus grazing 19 2 2.5 2 3 3 Capra hircus grazing 18 2 2.5 2.5 2.75 2.5 Capra hircus grazing 18 1 2 2 2 2 Average 1.85 2.6 2.5 2.75 2.80 Second goat experiment Capra hircus browsing 10 1.5 1.5 1 2 1 Capra hircus browsing 20 1.5 1.5 1 2 1 Capra hircus browsing 30 1.5 1.5 1 2 1 Capra hircus browsing 40 1 1 1 1 1 Average 1.28 1.50 1 1.87 1.31 Capra hircus grazing 10 0 2 2 2 1 Capra hircus grazing 20 1 2.5 2 3 2 Capra hircus grazing 30 1 3 3 3 2 Capra hircus grazing 40 1.5 3 3 3 2 Average 0.8 2.6 2.5 2.75 1.75 Species of Browsers Species # M-II M-III AB P B J P Okapia johnstoni 51222 0.5 1.5 2 1 1 Okapia johnstoni 51215 0 1 1 1 1 Okapia johnstoni 51229 0 1 1 1 1 Okapia johnstoni 51218 1 3 3 3 3.5 Okapia johnstoni 113805 1 1 2 2 2 Okapia johnstoni 51214 0 0 1.5 1.5 1 Okapia johnstoni 51225 1 1 1 1 2 Okapia johnstoni 51197 2 1 1 1 1 Okapia johnstoni 51227 1.5 1 1 1 2 Okapia johnstoni 51228 0.5 1.5 2 1 1 Okapia johnstoni 51219 1 1 1 1 1 Avg. 0.77 1.40 1.50 1.31 1.50 Giraffa camelopardalis 54123 2 1.5 1 2.5 1 Giraffa camelopardalis 53548 0 1 2 1 2 Giraffa camelopardalis 53546 2 1 1 2 2
194
Appendix 2 (continued) Species Species # M-II M-III AB P B J P Giraffa camelopardalis 82001 1 1 1 1 2 Giraffa camelopardalis 27753 1 1 1 1 1 Giraffa camelopardalis 24291 2 1.5 1 2 2 Giraffa camelopardalis 83548 0 1 1 1 2 Giraffa camelopardalis 242992 1 1 1 1 2 Giraffa camelopardalis 24290 - 1 - 1 - Giraffa camelopardalis 24293 4 2 2 2 2 Giraffa camelopardalis 16502 1 2 2 2 1 Giraffa camelopardalis 605051 0 1 1 1 1 Giraffa camelopardalis 165052 1 2 2 2 1 Giraffa camelopardalis 53549 1 1.5 1 2 1 Giraffa camelopardalis 83458 1.5 1.5 1 2 2.5 Avg. 1.25 1.33 1.28 1.56 1.60 Alces alces 3928 0.5 1.5 2 1 2 Alces alces 87098 1 1 1 1 1 Alces alces 122674 0 1 1 1 1 Alces alces 2191 2 1 1 1 2.5 Alces alces 2196 0.5 1 1 1 1 Alces alces 2192 2 1 1 1 2 Alces alces 2186 2 1 1 1 1.5 Alces alces 2193 0.5 2 2 2 2 Alces alces 2195 1.5 1.5 1 2 1.5 Alces alces 2187 2 1.5 1 2 2 Alces alces 2182 1.5 1 1 1 1.5 Alces alces 2200 0 1 1 1 1 Alces alces 2185 0.5 1.5 1 2 2 Grazers Avg 1.07 1.23 1.15 1.30 1.61 Ourebia ourebi 34764 2 3.5 3.5 3.5 3.5 Ourebia ourebi 27792 3 4 4 4 4 Ourebia ourebi 82051 2 4 4 4 4 Ourebia ourebi 34762 2 4 4 4 4 Ourebia ourebi 5419 2 4 4 4 4 Ourebia ourebi 54196 2 2.5 3 2 4 Ourebia ourebi 54194 2.5 2.5 2.5 3 Avg 2.16 3.50 3.57 3.42 3.57 Kobus ellipsiprymnus 80488 2 2 2 2 2 Kobus ellipsiprymnus 54198 4 4 4 4 4 Kobus ellipsiprymnus 53512 3 4 4 4 4 Kobus ellipsiprymnus 80489 2 4 4 4 3 Kobus ellipsiprymnus 8486 3 2 2 2 3 Kobus ellipsiprymnus 27773 2.5 2 2 2 3 Kobus ellipsiprymnus 27771 4 3 3 3 4 Kobus ellipsiprymnus 54201 4 4 4 4 4 195
Appendix 2 (continued) Species Species # M-II M-III AB P B J P Kobus ellipsiprymnus 27774 3 4 4 4 4 Kobus ellipsiprymnus 54200 2.5 2 2 2 4 Avg. 3 3.1 3.1 3.1 3.5 Connochaetes taurinus 3009 3 3 3 3 2 Connochaetes taurinus 3012 3 4 4 4 4 Connochaetes taurinus 3045 5 4 4 4 4 Connochaetes taurinus 3025 5 4 4 4 4 Connochaetes taurinus 3035 3 4 4 4 3.5 Connochaetes taurinus 3046 5 4 4 4 4 Connochaetes taurinus 3039 5 4 4 4 4 Connochaetes taurinus 3038 4 4 4 4 4 Connochaetes taurinus 3042 2 3 3 3 3 Connochaetes taurinus 3031 5 4 4 4 4 Connochaetes taurinus 3029 5 4 4 4 4 Connochaetes taurinus 3033 3 4 4 4 3.5 Avg. 4 3.8 3.8 3.8 3.6 Mixed feeders Cervus Canadensis 123172 0.5 1 1 1 1 Cervus Canadensis 40005 1 1 1 1 1 Cervus Canadensis 27406 0.5 1 1 1 1 Cervus Canadensis 120861 2 3.5 3.5 3.5 3 Avg. 1 1.6 1.6 1.6 1.5 Gazella granti 27662 4 4 4 4 Gazella granti 179196 1 2.5 3 2 2 Gazella granti 85151 3.5 2 1 3 4 Gazella granti 27793 1 4 4 4 4 Gazella granti 27791 1 1 1 2 Gazella granti 82152 2 2 2 2 Gazella granti 54150 2 2 2 2 2 Gazella granti 179199 2 2 2 2 2 Gazella granti 54157 3 2 1 3 Gazella granti 54146 3 3 3 3 Gazella granti 179270 2 2.5 2 3 3 Gazella granti 179198 1.5 1 2 2 Gazella granti 54151 3 4 4 4 4 Gazella granti 187831 3 2 2 2 3 Gazella granti 54148 2 4 Gazella granti 54149 1.5 2.5 2 3 Gazella granti 179201 3 2.5 2 3 2 Average 2.16 2.46 2.33 2.64 2.86 Mammalian Fossils Species Species # M-II M-III AB P B J P Progiraffa exigua 41662 1 1.25 1 1.5 1 196
Appendix 2 (continued) Species Species # M-II M-III AB P B J P Progiraffa exigua 41662 1 1.25 1 1.5 1 Progiraffa exigua 664 2 3 3 3 2 Progiraffa exigua 208 2 3 3 3 3 Progiraffa exigua 312 1.5 1.5 2 1 2 Progiraffa exigua 32173 1 1.25 1.5 1 1 Progiraffa exigua 5305 2 2 2 2 2 Avg. 1.5 1.89 1.92 1.85 1.71 Giraffokeryx punjabiensis AMNH19479 1 2 2 2 1 Giraffokeryx punjabiensis AMNH 95112 0 1 1 1 1 Giraffokeryx punjabiensis AMNH 406 0 1 1 1 1 Giraffokeryx punjabiensis AMNH 19475 2 1 1 1 1 Giraffokeryx punjabiensis Cast 62 1 1 1 1 1 Giraffokeryx punjabiensis 1023 2 1.25 1 1.5 1 Giraffokeryx punjabiensis 8217/1103 2.5 1.25 1 1.5 3 Giraffokeryx punjabiensis 30628 1.5 1.5 1 2 1.5 Giraffokeryx punjabiensis 30630 2 1.5 1.5 1.5 1.5 Giraffokeryx punjabiensis 30627 2 1.25 1 1.5 2 Giraffokeryx punjabiensis 41327 2 1.25 1 1.5 1 Giraffokeryx punjabiensis 40913A 1.5 1 1 1 1 Giraffokeryx punjabiensis 41328 2 1.25 1 1.5 2 Giraffokeryx punjabiensis 52360 2 1.25 1 1.5 1.5 Giraffokeryx punjabiensis 15136 Y452 1 1.75 1.5 2 1 Giraffokeryx punjabiensis 49576 Y 406 2 1 1 1 2 Giraffokeryx punjabiensis 49577 Y 406 2 1 1 1 1.5 Giraffokeryx punjabiensis 18089 Y 451 2 2.25 2 2.5 2.5 Giraffokeryx punjabiensis 23646 Y661 0 1 1 1 1 Giraffokeryx punjabiensis 23500 Y674 1.5 1 1 1 1 Giraffokeryx punjabiensis 27733 Y loc 729 1 1 1 1 1 Giraffokeryx punjabiensis 309889 loc 710 2 1 1 1 1 Giraffokeryx punjabiensis 1402 Y038 1 1.25 1 1.5 1 Giraffokeryx punjabiensis 32620 loc 752 2 1 1 1 1 Giraffokeryx punjabiensis 20142 Y061 1.5 1.5 1.5 1.5 1 Giraffokeryx punjabiensis 5694 Y017 2 1.25 1 1.5 2 Giraffokeryx punjabiensis 21926 Y504 2 1 1 1 1 Giraffokeryx punjabiensis 32578 Y500 2 1 1 1 1.5 Giraffokeryx punjabiensis 27666Y500 1.5 1.25 1 1.5 1 Giraffokeryx punjabiensis 23075Y500 2 1.25 1 1.5 1 Giraffokeryx punjabiensis 31702 Y503 1.5 1 1 1 2 Giraffokeryx punjabiensis 31703 Y503 1 1 1 1 2 Giraffokeryx punjabiensis 20583 Y640a 1 1 1 1 1 Giraffokeryx punjabiensis 23172 2 1 1 1 1.5 Giraffokeryx punjabiensis 24335 Y060 1 1.25 1 1.5 1 Giraffokeryx punjabiensis 20089Y060 1.5 1.5 1.5 1.5 1 197
Appendix 2 (continued) Species Species # M-II M-III AB P B J P Giraffokeryx punjabiensis 47142 Y641 0.5 1 1 1 1 Giraffokeryx punjabiensis 15205 Y038 0.5 1.25 1 1.5 1 Giraffokeryx punjabiensis 17160 Y496 1 1.25 1 1.5 1.5 Giraffokeryx punjabiensis 30774 Y496 2 1.25 1 1.5 1.5 Giraffokeryx punjabiensis 26648 Y 695 1.5 1 1 1 1 Giraffokeryx punjabiensis 27737 Y695 1 1 1 1.5 Giraffokeryx punjabiensis 20591 Y59 1 1.25 1 1.5 1 Giraffokeryx punjabiensis 19842 Y95 2 1.25 1 1.5 1 Giraffokeryx punjabiensis 12501 Y329 1 1.25 1 1.5 1 Giraffokeryx punjabiensis 24320 Y647 1 1.25 1 1.5 1 Giraffokeryx punjabiensis 24329 Y 647 2 1.5 1.5 1.5 1 Giraffokeryx punjabiensis 47141 Y647 2 1 1 1 1 Giraffokeryx punjabiensis 27860 Y647 2 1.25 1 1.5 2 Giraffokeryx punjabiensis 26466 Y059 1 1 1 1 1 Giraffokeryx punjabiensis 26465 Y059 2 1.25 1 1.5 1 Giraffokeryx punjabiensis 27668 Y498 2 1.25 1 1.5 1.5 Giraffokeryx punjabiensis 28113 Y705 2 1 1 1 1 Giraffokeryx punjabiensis 30667 1.5 1 1 1 1 Avg. 0.80 1.20 1.2 1.2 1? G. punjabiensis small 47323Y478 2 1 1 1 2 G. punjabiensis small 47324 Y 478 2 1.25 1 1.5 2 G. punjabiensis small 14778 2 1.25 1 1.5 1 Avg. 0.80 1.20 1.2 1.2 1.66 Giraffa sp. 1.5 1 1 1 1 1 Bramatherium small 48771 3 3 3 3 Bramatherium small 13201 1 1.25 2 3 1 Bramatherium small 12210 loc 311 1 1 1 1 1 Bramatherium small 6399 loc 311 1.5 1 2 1 Bramatherium small 49665 loc R 403 2.5 2 2 2 2 Bramatherium small 14395 2 2.75 2.5 2.5 1 Bramatherium small 9267 Y336 1 1 1 1 1 Bramatherium small 27662 Y507 1 1.25 1 1.5 1 Bramatherium small 4445 1.25 1 1.5 1 Bramatherium small 5361 Y076 1 1 1 1 1.5 Avg. 1.28 1.60 1.55 1.75 1.35 Bramatherium large 13553Y356 2 3 3 3 1 Bramatherium large 4920 Y196 2 4 4 4 3 Bramatherium large 53270 3 3.5 3 4 4 Bramatherium large 6002Y337 0 1 1 1 1 Bramatherium large 4238 Y142 1 1 1 1 2 Avg. 1.6 2.50 2.4 2.6 2.2 Bramatherium 11295 2 1.75 1.5 2 2 Bramatherium 16401loc 441 1.5 1.5 1.5 1.5 1 198
Appendix 2 (continued) Species Species # M-II M-III AB P B J P Bramatherium 18328 2 1.5 1 2 1.5 Bramatherium 15486 loc 458 2 1.75 1.5 2 2 Bramatherium 18412loc 605 2 1 1 1 1 Bramatherium 2798 loc Y019 2 1.25 1 1.5 2 Avg. 1.91 1.45 1.25 1.66 1.58 Asina gen. nov. sp. nov 15821Y517 0.5 1 1 1 1 Asina gen. nov. sp. nov 772 Y038 1 1.25 1 1.5 1 Asina gen. nov. sp. nov 15205 2 1.25 1 1.5 2 inclined slanted. Avg. 1.16 1.16 1 1.33 1.33 Paleotragus V55 inclined 1211 Y066 1 1 1 1 2 Pachyportax nagrii Cast 13 2 2 2 2 1 Pachyportax nagrii Cast 32 1.5 1 1 1 1 Pachyportax nagrii Cast 23 2 2 2 2 2 Pachyportax nagrii Cast 45 2-3 2.5 3 2 2 Avg 2 1.87 2 1.75 1.5 E. khauristanensis Cast 26 0-1 1 1 1 1 E. khauristanensis Cast 50 0-1 1.5 1 2 1 Avg.mesoband2 1 1.25 1 1.5 1 Gazella lydekkeri Cast 20 1-2 2.5 2 3 1 Gazella lydekkeri GCUPC 1008/07 1 1 1 1 1 Gazella lydekkeri Cast 78 2 2.5 2 3 2 Gazella lydekkeri GCUPC 1089/07 2 2.5 3 2 2 Gazella lydekkeri GCUPC 1079/07 2 2.5 2 3 2 Gazella lydekkeri GCUPC 1269/09 2 2 2 2 2 Average 1.75 2.16 2 2.33 1.66 Tragoportax sp. Cast 48 0-1 2 2 2 1.5 Tragoportax sp. Cast 37 1 2 2 2 1.5 Tragoportax sp. Cast 46 1.5 2 2 2 1 Tragoportax sp. Cast 70 1.5 1.75 2.5 2 1.5 Tragoportax sp. Cast 73 0-1 1 1 1 2 Average 1 1.34 1.43 1.37 1.5 Dorcatherium majus Cast 47 0.5 2 2 2 1 Dorcatherium majus Cast 77 0.5 1 1 1 1 Dorcatherium majus Cast 24 0.5 1 1 1 1 Dorcatherium majus Cast 67 2 2 2 2 1 Dorcatherium majus Cast 16 1 1.5 1 2 1 Dorcatherium majus Cast 2 2 2 2 2 Average 1.08 1.58 1.5 1.66 1.16 Dorcatherium minus Cast 76 2 1.5 2 1 1 Merycopotamus nanus Cast 41 1 1 1 1 1 Merycopotamus nanus Cast 12 2 2 2 2 2 Average 1.5 1.5 1.5 1.5 1.5 Miotragocerus gluten Cast 27 2 2 2 2 2 199
Appendix 2 (continued) Species Species # M-II M-III AB P B J P Pachyportax latidens Cast 35 2 3 3 3 3 Selenoportax lydekkeri Cast 68 2.5 2.5 2 3 2 Selenoportax lydekkeri Cast 15 3 3 3 3 3 Average 2.75 2.75 2.5 3 2.5 Selenoportax vexillarius Cast 45 3 3 3 3 2.5 Selenoportax vexillarius Cast 69 4 3 3 3 2.5 Selenoportax vexillarius Cast 65 4 3 3 3 3 Selenoportax vexillarius Cast 64 2 2.5 2 3 2 Average 3.25 2.87 2.75 3 2.5 Hipparion sp. Cast 34 6 4 4 4 4 Hipparion sp. GCUPC 1126 4 4 4 4 4 Hipparion sp. GCUPC 1127 3 3 3 3 3 Hipparion sp. GCUPC 1431 4 4 4 4 4 Hipparion sp. GCUPC 1118 4 3.5 4 3 3 Hipparion sp. GCUPC 1132 3 3 3 3 2.5 Hipparion sp. GCUPC 1107 4 3.5 3 4 4 Hipparion sp. GCUPC 1102 4 4 4 4 4 Hipparion sp. GCUPC 1121 4.5 4 4 4 4 Average 4.9 3.85 3.85 3.85 3.78 Fossils of giraffid from China Samotherium boissieri 481 2.5 3 3 3 4 Samotherium boissieri 56 A0086 2 1.25 1.5 1 2 Samotherium boissieri 29X026 3 3 3 3 4 Samotherium boissieri 202D0078 2 1.5 1 2 1.5 Samotherium boissieri 1X1240 0.5 1.5 2 1 1 Samotherium boissieri 550 2 3 3 3 2 Samotherium boissieri NS 90HMV1573 2 - - - - Samotherium boissieri NS 13 0944 1 1.5 2 1 1 Average 1.9 2.10 2.21 2.10 2.21 Samotherium sp. 25 M0274 2 - - - - Samotherium sp. Qiui number 1 - 3 2 4 3 Samotherium sinense 1393-32610 4 4 4 4 4 Samotherium sinense 120949 - 3 3 3 2 Samotherium sinense ZD 0701 Average 3.33 3.33 3.33 2.6 Honanotherium schlosseri 60 M0336 2 1.5 2 1 2 Honanotherium schlosseri 41C0053 1.5 2.75 2.50 3 3.50 Honanotherium schlosseri 20146 2 3 3 3 4 Honanotherium schlosseri 20359 2 1.5 2 1 1 Honanotherium schlosseri 53? C0089 2 1.5 2 1 1 Honanotherium schlosseri 57A0089 1 2 2 2 3.5 Average 1.9 2.17 2.35 2 2.64 Palaoetragus coelophrys 206M0332 2 2 2 2 2 200
Appendix 2 (continued) Species Species # M- M-III AB P B J P II Palaoetragus coelophrys 3161 2.5 4 4 4 4 Palaoetragus coelophrys 61C0048 2 2.5 2 3 2 Palaoetragus coelophrys 40834 2 3 3 3 3 Palaoetragus coelophrys 35 C0085 2 3 3 3 1 Palaoetragus coelophrys 23480 2 1.5 2 1 4 Palaoetragus coelophrys 300953 2 2 2 2 2.75 Palaoetragus coelophrys NS 10 - - - - - Average 2.1 2.66 2.66 2.66 2.86 Paleotragus rouenii 22 (12) 2.5 2.5 2 3 2 Paleotragus rouenii NS3 M0018 1 - - - - Average 1.7 2.5 2 3 2 Schasitherium tafeli A025 2 4 4 4 4 Schasitherium tafeli 46162171 2 3 3 3 2 old Schasitherium tafeli 441740 0.5 1.5 2 1 1 Schasitherium tafeli 20037 - 2.5 3 2 3.5 Schasitherium tafeli 49X1238 3 2 2 2 2 Schasitherium tafeli 201M0335 - 1.5 2 1 1 Schasitherium tafeli 90945 - 1 1 1 1 Schasitherium tafeli 2529 - 1 3 3 3 Schasitherium tafeli NS 54A0256 2 3 3 3 3 Schasitherium tafeli NS55 HMV1572 2 - - - - Average 1.9 2.16 2.55 2.22 2.27 Key to symbols; M-II=mesowear type II, M-III=mesowear type III, AB=anterior band, PB=posterior band, JP= junction point. Appendix 3; Microwear data for Siwalik ungulates. Propotamochoerus hysudricus Sample# Locality Age AP LP G AS Cr.S FS CS MS Y17578 Y311 10.0 34 1 1 20.5 1 1 Y22080 Y311 10.0 31.5 1 0 7 1 1 Y6226 Y228 9.50 26 1 1 17.5 1 1 Y14044 Y262 9.5 23 1 0 27.5 1 1 Y17088 Y262 9.5 49 1 1 13.5 1 1 Y6011 Y341 9.3 35 1 1 11 1 1 Y14501 Y269 9.3 23 1 1 19 1 1 Y14502 Y269 9.3 28 1 0 19 1 1 Y41368 Y269 9.3 23.5 1 1 14 1 1 Y10224 Y309 9.3 37 1 1 15 1 1 Y13587 Y227 9.3 21.5 1 1 8 1 1 Y6137 Y221 9.3 19 1 1 18 1 1 Y16132 Y310 9.3 22.5 1 0 19.5 1 1 Y10998 Y314 9.2 27.5 1 1 20.5 1 1 201
Appendix 3 (continued) Sample# Locality Age AP LP G AS Cr.S FS CS MS Y11007 Y314 9.2 26.5 1 1 19.5 1 1 Y5112 Y211 9.2 27 1 0 11.5 1 1 Y9355 Y260 9.2 36 1 1 20 1 1 Y4554 Y182 9.2 55 1 0 19 1 1 Y7654 Y182 9.2 47 1 1 16 1 1 Y13678 Y367 8.9 58.5 1 1 22.5 1 1 Y4599 Y193 8.9 41.5 1 1 19 1 1 Y17968 Y604 8.5 20.5 1 1 17 1 1 Y4813 Y203 8.5 12 1 0 19 1 1 Y457 Y024 8.1 24.5 1 1 20.5 1 1 Y49515 Y599 8.0 26 1 0 19.5 1 1 Y50880 Y949 7.8 34.5 1 0 21.5 1 1 Y49840 Y906 7.8 39 1 1 8.5 1 1 Y49844 Y906 7.8 28 1 1 23 1 1 Y18124 Y452 7.2 22 1 0 15.5 1 1 Y14045 Y370 7.1 11.5 1 1 13 1 1 Y49816 Y916 7.0 32.5 1 0 14.5 1 1 Y85 YO10 7.0 42.5 1 1 23 1 1 Y2807 Y105 7.0 42 0 0 18.5 1 1 Y19698 D010 7.0 31.5 1 0 18.5 1 1 Y19748 L37 6.5 30.5 1 0 24 1 1 Y19803 D012 6.5 24.5 1 1 20.5 1 1 Y51575 Y539 6.5 27.5 1 1 19 1 1 Y51848 Y269 6.5 27 1 0 20 1 1 Y52623 Y960 6.5 37.5 1 1 20 1 1 Y52855 O84 6.5 30.5 1 1 7.5 1 1 D3 D005 6.5 24.5 1 1 11 1 1 CST 28 PDR 6.0 17 - - 25.5 - - - - CST 40 PDR 6.0 09 - - 30 - - - - CST 39 HNT15 6.0 15 - - 29 - - - - CST 25 MM9 6 08 - - 32 - - - - CST 33 MM10 5 19.5 - - 25.5 - - - - CST 11 HNT-17 5 09 - - 20.5 - - - - AVG 28.46 18.59 SD 10.2 4.7 % 97.6 61.0 100.0 9.8 7.3 82.9 MI=0.65 Hippopotamodon sivalense Y18261 Y454 10.4 29 1 0 25.5 1 1 Y18264 Y454 10.4 37 1 1 20.5 1 1 Y18276 Y454 10.4 28 0 0 10.5 1 1 Y18280 Y454 10.4 42 1 1 16 1 1 202
Appendix 3 (continued) Sample# Locality Age AP LP G AS Cr. S FS CS MS D36 D27 10.4 20 1 1 11.5 1 1 Y53009 Y450 10.2 13.5 0 0 21 1 1 Y41401 Y311 10.0 36 1 1 18 1 1 Y13972 Y251 10.0 56 1 1 14 1 1 Y16049 Y251 10.0 28.5 1 0 23 1 1 Y46568 Y251 10.0 23.5 1 1 16 1 1 Y12861 Y309 9.3 28 1 1 21.5 1 1 Y4984 Y159 9.3 30.5 1 0 28 1 1 Y5276 Y211 9.2 11 1 0 20.5 1 1 Y13524 Y239 9.2 25.5 0 0 18 1 1 Y14112 Y260 9.2 31 1 1 15 1 1 Y27915 Y260 9.2 43 1 1 23.5 1 1 Y16085 Y182 9.2 28.5 1 1 21 1 1 Y17937 KL12 8.5 12.5 0 0 17 1 1 Y4086 Y174 8.3 26 1 1 16 1 1 Y330 YO17 8.0 29.5 0 0 16.5 1 1 Y50019 Y917 7.8 27 1 0 19.5 1 1 Y50457 Y947 7.8 9 0 0 17.5 1 1 Y50134 Y932 7.2 30 1 0 22 1 1 CST-30 HNT 7.00 13 - - 26.5 - - - - AVG 27.41 19.10 STDEV 10.8 4.3 MI=0.69 %age 73.9 39.1 100 4.3 13.0 82.6 Dorcatherium majus D482 D027 10.4 39 1 1 17 1 1 Y15126 Y450 10.1 22.5 1 1 14.5 0 1 Y16847 Y450 10.1 21 1 1 17 1 1 Y 18007 Y450 10.1 17.5 1 0 18.5 1 1 Y50639 Y450 10.1 24 1 1 21 0 1 Y50647 Y450 10.1 31 1 1 18.5 0 1 Y11368 Y311 10.0 19.5 1 0 18.5 1 1 Y11391 Y311 10.0 30.5 1 0 31 1 1 Y17707 Y311 10.0 19.5 0 0 25 1 1 Y17748 Y311 10.0 21 0 0 15 1 1 Y19949 Y311 10.0 28.5 1 0 13 0 1 Y20169 Y311 10.0 12 0 1 20 1 1 Y20552 Y311 10.0 27 1 1 17.5 1 1 Y22082 Y311 10.0 23 1 0 10 0 1 Y22083 Y311 10.0 17 1 0 22.5 1 1 Y27827 Y311 10.0 30.5 1 0 15 1 1 Y41417 Y311 10.0 33 1 0 15 1 1 Y41418 Y311 10.0 33 1 1 11.5 0 1 203
Appendix 3 (continued) Sample# Locality Age AP LP G AS Cr.S FS CS MS Y47503 Y311 10.0 30.5 1 0 20.5 1 1 Y47533 Y311 10.0 25.5 1 0 23.5 1 1 Y47604 Y311 10.0 11.5 1 0 22 1 1 Y47671 Y311 10.0 27.5 1 0 14.5 1 1 Y47680 Y311 10.0 22.5 0 0 11.5 1 1 Y5562 Y310 9.3 46 1 0 20.5 1 1 Y49530 Y888 8.0 27 1 1 13 1 1 Y49555 Y539 8.0 44.5 1 0 16 0 1 Y16236 Y540 8.0 30 1 0 22 1 0 Y50441 Y947 7.8 30 1 0 12 0 1 Y52556 Y941 7.3 38.5 1 0 16 0 1 Y19707 7.0 56 1 0 11 1 1 Y52833 Y981 7.0 13.5 0 0 22.5 1 1 Y52856 LO84 7.0 37.0 0 0 9.0 0 1 CST 47 HNT 6.5 11.00 - - 19.50 - - - - CST 42 HNT 6.5 7.500 - - 14.50 - - - - CST 16 PDR 6.0 12.00 - - 21.50 - - - - CST 24 PDR 6.0 17.50 - - 17.50 - - - - AVG 26.04 - - 17.43 - - - - STDEV 10.0 4.9 MI=0.66 % 81.3 28.1 68.8 15.6 3.1 81.3 Selenoportax vexillarius CST 78 HNT 7 9.50 - - 18 - - - - CST 50 PDR 6 10 - - 22 - - - - CST 64 PDR 6 17 - - 29.5 - - - - CST 68 HNT 6 22 - - 13.5 - - - - CST 65 MM1 6 26 - - 09 - - - - CST 69 MM1 6 20 - - 13.5 - - - - AVG. 17.41 17.58 MI=1.00 STDEV 5.90 6.19 Pachyportax latidens CST35 HNT 8 24 - - 34 - - - - CST17 MM3 8 16 - - 18 - - - - CST19 PDR 6 13 - - 33 - - - - CST66 PDR 6 22.5 - - 24.5 - - - - AVG. 18.87 - - 27.37 - - - - MI=1.45 STDEV 8.18 6.55 Tragoportax sp. CST46 MM4 8 15 23 CST37 MM2 7.5 14 28.50 CST70 MM2 7.5 26 18 CST67 PDR 6 12 23.50 204
Appendix 3 (continued) Sample# Locality Age AP LP G AS Cr.S FS CS MS CST74 PDR 6 18 18.50 CST73 MM1 6 17 24 AVG. 17 22.58 MI=1.32 STDEV 4.47 3.55 Miotragocerus gluten CST27 7.5 12.5 - - 18 - - - - MI=1.44 Merycopotamus nanus CST12 MM4 8 14 - - 25.50 - - - - CST41 MM3 8 10 - - 17 - - - - AVG. 12.00 21.25 STDEV 2 4.25 MI=1.77 Hipparion sp. CST 34 HNT 7.5 09 - - 25.5 - - - - MI=2.83 Brachypotherium fatejhangense CST 61 MM4 8 32.50 11 MI=0.33 Brachypotherium perimense CST 62 MM2 7.5 11 - - 22 - - - - MI=2 Key to Microwear Symbols (headings) “AP”= Average Pits; “AS” Average Scratches; “CS” coarse scratches; LPs= large pits; “G”= gouges; “FS”= fine scratches; “Cr. S”= cross scratches; “MS”= mixed scratches. Microwear characters include continuous variables (number of pits and number of scratches) and categorical variables (large pits, gouges and cross scratches). The categorical variables include presence/absence of at least four of each of large pits, gouges and cross scratches. In each of the characters (both continuous and categorical), presence of character is indicated by “1” whereas no is represented by “0”. The texture of scratches is characterized by predominantly coarse, predominantly fine and mixed scratches (MS). The texture assigned to each specimen is represented by “1”, CST= cast. Studied specimens. Referred data were taken from Nelson (2003).
205
Appendix 4; Comparison of microwear data by two workers (NS magnification vs. MT magnification) Goat Experiment NS Magnification Grazers Cast # Tooth MS AP AS 01 Upper 1 30 26 10 Lower 3 28 13.5 4 Upper 1 20.5 20 5 Lower 1 15 26 3 Upper 2 24.5 29.5 6 Upper 1 29 13 31 Lower 33.5 23.5 7 Upper 0 15.50 24 2 Lower 1 37 23 MI=0.89 Aerage 25.88 23.16 MT Magnification 01 Upper 1 14 13 10 Lower 3 14 07 4 Upper 1 10 11 5 Lower 1 08 12 3 Upper 2 12 15 6 Upper 1 15 07 31 Lower 16 12 7 Upper 0 08 12 2 Lower 1 18 12 MI=0.84 Average 1.37 13.22 11.22 Browsers Cast # Tooth MS AP AS NS Magnification 55 Upper 0 16 27 54 Lower 2-3 28 29 52 Upper 2 32.5 23 59 Lower 2 30 26.5 57 Upper 0-1 14.5 29 9 Lower 2 48 16.5 51 Upper 0-1 32.5 28 58 Lower 1-2 26.5 23 56 Upper 2 24.5 32 53 Lower 0-1 17.5 23.5 MI= 0.95 Average 1.37 27 25.75 Browsers 55 Upper 0 8 13 MT Magnification 54 Lower 2-3 14 15 52 Upper 2 16 12 59 Lower 2 15 13 57 Upper 0-1 07 15 9 Lower 2 24 09 51 Upper 0-1 16 14 206
Appendix 4 (continued) Cast # Tooth MS AP AS 58 Lower 1-2 13 12 56 Upper 2 12 16 53 Lower 0-1 9 12 MI=0.97 Average 1.3 13.4 13.10 Propotamochoerus hysudricus NS Magnification CST 28 PDR 6.0 25.5 17 CST 40 PDR 6.0 30 09 CST 39 HNT15 6.0 29 15 CST 25 MM9 6.0 32 08 CST 33 MM10 5.0 25.5 19.5 CST 11 HNT-17 5.0 20.5 09 Microwear Index =0.47 27.08 12.91 MT Magnification CST 28 PDR 6.0 13 09 CST 40 PDR 6.0 14 05 CST 39 HNT15 6.0 15 08 CST 25 MM9 6.0 17 05 CST 33 MM10 5.0 12 10 CST 11 HNT-17 5.0 10.5 04 Microwear Index = 0.5 13.58 6.83 H. sivalense CST-30 HNT 7.00 26.5 13 NS Magnification MI=0.49 MT Magnification CST-30 HNT 7.00 13.5 7 MI=0.51 Dorcatherium majus CST 47 HNT 6.5 11.00 19.50 NS Magnification CST 42 HNT 6.5 7.500 14.50 CST 16 PDR 6.0 12.00 21.50 CST 24 PDR 6.0 17.50 17.50 MI= 1.53 Avg. 11.87 18.25 Dorcatherium majus CST 47 HNT 6.5 06 10 MT Magnification CST 42 HNT 6.5 5 7 CST 16 PDR 6.0 6 12 CST 24 PDR 6.0 10 10.5 MI= 1.46 Avg. 6.75 9.87 Selenoportax vexillarius NS Magnification CST 78 HNT 7 9.50 18 CST 50 PDR 6 10 22 CST 64 PDR 6 17 29.5 CST 68 HNT 6 22 13.5 CST 65 MM1 6 26 09 CST 69 MM1 6 20 13.5 MI=1.00 AVG. 17.41 17.58
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Appendix 4 (continued) Cast # Tooth MS AP AS STDEV 5.90 6.19 MT Magnification CST 78 HNT 7 4.5 8.5 CST 50 PDR 6 6 11.5 CST 64 PDR 6 9 14 CST 68 HNT 6 10 6 CST 65 MM1 6 13 4 CST 69 MM1 6 10 7 MI=1.03 AVG. 8.25 8.50 Pachyportax latidens NS Magnification CST35 HNT 8 24 34 CST17 MM3 8 16 18 CST19 PDR 6 13 33 CST66 PDR 6 22.5 24.5 MI= 1.45 AVG. 18.87 27.37 MT Magnification CST35 HNT 8 11.5 16 CST17 MM3 8 8 10 CST19 PDR 6 6 17 CST66 PDR 6 11 12 AVG. 9.12 13.75 MI=1.50 Tragoportax sp. NS Magnification CST46 MM4 8 15 23 CST37 MM2 7.5 14 28.50 CST70 MM2 7.5 26 18 CST67 PDR 6 12 23.50 CST74 PDR 6 18 18.50 CST73 MM1 6 17 24 MI= 1.32 AVG. 17 22.58 Tragoportax sp. MT Magnification CST46 MM4 8 7 12 CST37 MM2 7.5 6.5 14 CST70 MM2 7.5 13 10 CST67 PDR 6 9 12 CST74 PDR 6 11 10 CST73 MM1 6 8 12 MI=1.28 Avg. 9.08 11.66 Miotragocerus gluten NS Magnification CST27 7.5 12.5 18 MI=1.44 CST 27 7.5 6.5 9 MT Magnification MI=1.38 Merycopotamus nanus NS Magnification CST12 MM4 8 14 25.50 208
Appendix 4 (continued) Magnification Sample# Locality Age AP AS CST41 MM3 8 10 17 MI=1.77 AVG. 12.00 21.25 MT Magnification CST12 MM4 8 7.5 12 CST41 MM3 8 6 8.5 AVG. 6.75 10.25 MI=1.51 Hipparion sp. NS Magnification CST 34 HNT 7.5 09 25.5 MI=2.83 MT Magnification CST 34 HNT 7.5 5.0 13.00 MI=2.60 Brachypotherium fatejhangese NS Magnification CST 61 MM4 8 32.50 11 MI=0.33
MT Magnification CST 61 MM4 8 16 6.00 MI=0.37 Brachypotherium perimense NS Magnification CST 62 MM2 7.5 11 22 MI=2 MT Magnification CST 62 MM2 7.5 6.00 11.5 MI=1.91 Giraffokeryx sp. CST 16 47.5 16.5 CST 72 27.0 25.1 Avg. 37.25 20.8 NS Magnification MI=0.55 Giraffokeryx sp. CST 16 22.5 7.5 CST 72 13.0 14 Avg. 17.75 10.7 MT Magnification MI=0.60 Key to Sybmols; NS=Nikos Solounias, MT=Muhammad Tariq
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Appendix 5; Ecomorphic and biostratigraphic data of Siwalik anthracotheres, caprini, alcephini, doliochoerini, hippopotamidae, camilidae, chalicotherinae and proboscideans.
Taxon Biostratigraphic HI B.W Diet AI ranges (Kg) IFA ILA Anthracotheriidae Anthracotherium punjabiensis 14.00 12.50 - 50-250 SB R Microbunodon punjabiensis 13.80 8.40 - - SB R Anthracotheriinae Microbunodon milaensis sp.nov 10.30 9.20 - - SB R Microbunodon silistrensis 17.00 11.50 - - SB R Hemimeryx sp. 14.0 3.6 - 50-250 SB R Bothriodontinae Merycopotamus dissimilis older 13.90 8.50 - - MFC C Merycopotamus dissimilis 8.00 3.30 - - MFO C younger Merycopotamus medioximus 10.40 8.60 - SB C Merycopotamus nanus 13.90 11.30 - - SB A Caprini - Caprotragus potwaricus - - 20-100 MFC R Sivacapra sivalense - - 20-100 MFO C Alcephini - - Demalops palaindicus - - 125- MFO C 342 Doliochoerinae Schizochoerus gandakasensis 11.20 8.10 - 8-50 SB R Hippopotamidae Hexaprotodon sivalensis 6.10 2.20 - 235 FG R Camilidae Camelus sivalensis 3.50 0.50 - 50-250 MFO R Chalicotherini Chalicotherium salinum 12.9 8.00 - 250- BB R 1000 Proboscidea Elephantiidae-Elephantiinae Elephas planifrons 3.30 0.50 - 6000 GG C Elephas promaximus 2.50 0.50 - 6000 GG R Elephas hysudricus 3.50 0.50 - <6000 GG C Choerolophinae Synconolophus sp. 10.00 6.50 - HB C Synconolophus ptychodus 16.00 12.50 - HB C Stegotetrabelodontinae Stegotetrabelodon sp. 10.00 7.00 - >2160 BB Deinotheriidae 210
Appendix 5 (continued) Taxon Biostratigraphic HI B.W Diet AI ranges (Kg) IFA ILA Deinothriinae Deinotherium indicum 13.50 7.50 - >1000 BB R Deinotherium pentapotamie 18.0 6.90 >1000 BB C Prodeinotherium sp. 20.00 13.00 - - BB R Stegodon bombiferons 10.1 1.50 - 5000- MFO C 6000 Mammutidae Stegodontinae Stegodon sp. 2.50 0.50 - <6000 MFO R Stegodon insignis 3.50 0.50 - <6000 MFO R Anancinae Anancus osborni 5.50 2.50 - >5400 MFO C Anancus properimensis 13.50 11.00 - 4000 MFC R Pentalophodon sivalensis 2.50 0.50 - 6000 GG C Stegolophodontinae Stegolophodon cautleyi 14.00 4.50 - >1000 BB- R MFO Stegolophodon stegodontoides 13.50 4.20 - >1000 BB- C MFO Gomphotheriidae Tetralophodontinae Tetralophodon falconeri 13.00 8.00 - 1080 SB C Tetralophodon punjabiensis 10.00 2.50 - >1080 SB C Gomphotheriinae Gomphotherium sp. 18.00 13.00 - 2160 SB C Key to symbols “HB”= high level browser; “BB”= regular/unspecialized browser; “SB”= selective browser; “O”= omnivore “MFC”= mixed feeder in closed habitat; “MFO”= mixed feeder in open habitat; “FG”= fresh grass grazers; “GG”= dry grass grazer; AI=abundance index; C=common; A=abundant; R= rare, IFA= inferred first appearance, ILA= inferred last appearance. Estimated body weight/mass are from Vrba (1980); Flynn et al. (1995); Palmer (1999); Barry et al. (2012).
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Appendix 6; Faunal list of Primates, Lagomorphs and Carnivores from Siwaliks.
Taxon Biostratigraphic ranges H B.W Diet AI IFA (Ma) ILA (Ma) I (Kg) Primates IFA ILA Sivapithecus sp. 13.50 8.40 - SB C Sivaladapis palaindicus 13.80 11.00 - SB R Sivapithecus parvada 10.00 9.80 - SB R Sivapithecus (?) simonsi R Lagomorpha Caprolagus sivalensis Carnivores recorded from Siwaliks of Pakistan Carnivora Felidae Pachycrocuta brevirostris Lutra palaindica Amphicyon sp. (Chinji) Enhydriodon sivalensis Felidae small sp. (14-6.6 Ma) Vishnuictis durandi Eomellivora sp. (14-11.2 Ma) Hyaenictis bosei Vishnufelis sp. (Chinji) Crocuta colvini Viverra sp. (13.7-2.5 Ma) Crocuta sivalensis Crocuta feline Magnetereon falconeri Percrocutidae Magnetereon palaindicus Percrocuta carnifex (14-10.5 Ma) Felis subhimalayana Dissopalis carnifex (Chinji) Sivapanthera potens (=Sivafelis potens) Panthera sp. Sivapanthera brachygnathus (=Sivafelis brachygnathus) Procyonidae Melursus theobaldi Sivanasua himalayensis (Nagri-DP? Sinictis lydekkeri Sivanasua nagrii (NG) Mellivora sivalensis Mustelidae Sivaonyx bathygnathus (NG-DP) Enhydriodon falconeri (Ng-Dp) Viverridae Vishnuictis hariensis (NG-DP) Hynidae Crocuta gigantea (NG-Dp) Crocuta mordax/tatroti (NG-DP) Lycyaena macrostoma (NG-DP) Ictitherium nagrii (NG-DP) Felidae Vinayakia intermedia (NG-DP) Canis pinjorensis (Pinjor) Canis cautleyi Protocyon curvipalatus (Sivacyon curvipalatus) Agriotherium sivalense 212
Appendix 7; Raw mesowear data of Giraffokery punjabiensis, Tragoportax sp., selenoportax sp., Gazella lydekkeri. Cusp relief is categorized as high (H) or low (L) based on how high the projection of cusps above the inter-cusp valley. Catalogue No. Tooth Position MESOWEAR-I MESOWEAR-II Cusp relief Cusp shape MS Giraffokeryx punjabiensis GCUPC 1329/11 M2 H S 0-1 GCUPC 1335/11 M2 H S 0-1 GCUPC 1305/11 M2 H S 0-1 GCUPC 1276/10 M2 H S 0-1 GCUPC 1272/10 M2 H S 0-1 GCUPC 491/02 M2 H R 2-3 GCUPC 1336/11 M2 H R 2-3 GCUPC 1142/09 M2 H R 2-3 GCUPC 1166/09 M2 H R 2-3 GCUPC 490/02 M2 H R 2-3 GCUPC 1320/11 M2 H S 0-1 GCUPC 1138/09 M2 H R 2-3 GCUPC 1325/11 M2 H S 0-1 GCUPC 1174/09 M2 H S 0-1 GCUPC 1172/09 M2 H S 0-1 GCUPC 1135/09 M2 H S 0-1 GCUPC 1275/10 M2 H S 0-1 Tragoportax sp. GCUPC 1297/09 M2 H S 0-1 GCUPC 1358/10 M2 H S 0-1 GCUPC 1321/10 M2 H S 0-1 GCUPC 1092/08 M2 H S 0-1 GCUPC 1319/09 M2 H S 0-1 GCUPC 1235/09 M2 H R 2-3 GCUPC 1239/09 M2 H R 2-3 Pachyportax latidens GCUPC 1322/09 M2 H S 0-1 GCUPC 1065/08 M2 H R 2-3 GCUPC 1009/07 M2 H S 0-1 GCUPC 1019/07 M2 H S 0-1 GCUPC 1016/07 M2 H R 2-3 GCUPC 1028/09 M2 H S 0-1 GCUPC 1262/09 M2 H S 0-1 GCUPC 1264/09 M2 H S 0-1 Selenoportax sp. GCUPC 1060/07 M2 H S 0-1 213
Appendix 7 (continued) Catalogue No. Tooth Position MESOWEAR-I MESOWEAR-II Cusp relief Cusp shape MS GCUPC 1282/09 M2 H S 0-1 GCUPC 436/02 M2 H R 2-3 GCUPC 438/02 M2 H S 0-1 GCUPC 493/02 M2 H S 0-1 PUPC 83/90 M2 H R 2-3 PUPC 85/06 M2 H S 0-1 PUPC 85/37 M2 H S 0-1 PUPC 86/201 M2 H S 0-1 PUPC 87/195 M2 H S 0-1 PUPC 85/37 M2 H R 2-3 GCUPC 1241/09 M2 L R 4-5 Gazella lydekkeri GCUPC 1060/07 M2 H S 0-1 PUPC 95/01 M2 H S 0-1 GCUPC 436/02 M2 L R 4-5 GCUPC 438/02 M2 H S 0-1 GCUPC 493/02 M2 H S 0-1 PUPC 83/90 M2 L R 4-5 PUPC 85/06 M2 H S 0-1 PUPC 85/37 M2 L R 0-1 PUPC 86/201 M2 H S 0-1 PUPC 87/195 M2 H S 0-1 PUPC 85/82 M2 H R 2-3 PUPC 00/101 M2 L R 4-5 Key to symbol: MS=mesowear scale
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Appendix 8; Hypsodonrty measurements of some artiodactyles incorporated for mesowear analysis Species Catalogue No. Tooth H W HI Position Giraffokeryx GCUPC 959/08 lm3 21.16 17.45 1.21 punjabiensis GCUPC 1286/10 rm3 18.00 15 1.20 GCUPC 1308/11 lm3 27.00 22.75 1.67 GCUPC 1242/10 lm3 21.00 17.00 1.23 GCUPC 1243/10 rm3 25. 16 18.00 1.38 GCUPC 1240/10 lm3 22. 12 17.50 1.25 GCUPC 1249/10 rm3 24. 16 19.00 1.26 Mean HI= 1.31±0.06 1.31±0.06 Tragoportax sp. GCUPC 1392/10 m3 13.23 11 1.20 GCUP C 1361/10 m3 13.15 10 1.31 GCUPC 1003/07 m3 13.55 11 1.23 GCUPC1330/09 m3 18.37 12.69 1.44 Mean HI= 1.29±0.05 1.29±0.05 Selenoportax sp. GCUPC 1326/09 m3 31 12.79 2.42 GCUPC 1386/10 m3 27 14.75 1.83 GCUPC 1075/07 m3 25.76 12.68 2.03 GCUPC 1057/07 m3 28.15 15.00 1.87 Mean HI=2.03±0.13 2.03±0.13 Gazella lydekkeri GCUPC 1098/07 m3 35.40 18.42 1.92 GCUPC 1388/10 m3 35.36 18.22 1.94 GCUPC 1004/07 m3 28.81 15.81 1.82 GCUPC 1295/09 m3 28.57 16.04 1.78 GCUPC1383/10 m3 28.30 12.69 2.23 GCUPC 1098/07 m3 46.25 18.76 2.46 Mean HI=2.18±0.10 2.18±0.10
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Appendix 9; Biostratigraphic ranges of fossils species of ungulates from Pakistan Siwaliks. Family/ Species Biostratigraphic Ranges N O SubFamily/Tribe ILA LO FO IFA I (Ma) (Ma) (Ma) (Ma) Stegodontidae Stegodon bombiferons - 3.5 10.1 - - Stegodontidae Stegodon insignis - 0.50 3.5 - - Stegodontidae Stegodon sp. - 0.78 2.5 - - Elephantidae Elephas planifrons - 1.50 3.5 - - Elephantidae Elephas proximuns - 0.50 2.5 - - Elephantidae Elephas hysudricus - 0.50 1.7 - - Gomphothriidae Synconolophus corrugatus 3.5 10.3 Gomphothriidae Synconolophus ptychodus - 12.5 16.5 - - Gamphotheriidae Gomphotherium sp. - 13.0 18.3 - -
Dinotheriidae Deinotherium indicum - 7.5 13.5 - - Dinotheriidae Deinotherium pentapotamie - 6.9 18.3 - - Dinotheriidae Prodeinotherium sp. - 13.0 20.0 - - Anancinae Tetralophodon sp. - 3.5 13.5 - - Anancinae Anancus properimensis 11.2 13.5 Anancinae Anancus osborni - 2.5 5.5 - -
Anancinae Pentalophodon sivalensis - 0.5 2.5 - -
Stegolophodontinae Stegolophodon cautelyei - 4.50 14.0 - -
Stegolophodontinae Stegolophodon stegodontoides - 4.2 13.5 - -
Stegotetralophodont Stegotetralophodon sp. - 7.0 10.5 - - -inae Choerolophinae Synconolophus ptychodus - 13.0 18.3 - - Choerolophinae Choerolophodon corrugates - 6.5 13.5 - - Tayassuidae Pecarichoerus orientalis - 11.0 13.5 - - Suidae Bunolistriodon gaptae 15.0 18.3 Suidae Palaeochoerus pascoei - 14.0 20.0 - - 216
Appendix 9 (continued) Family/ Species Biostratigraphic Ranges N O SubFamily/Tribe ILA LO FO IFA I (Ma) (Ma) (Ma) (Ma) Suidae Palaeochoerus perimense - 11.0 14.2 - -
Suidae Listriodon pentapotamie - 10.3 14.0 - -
Suidae Lophokeryx nagrii - 9.50 11.0 - -
Suidae Hyotherium chisholmi - 11.0 13.2 - -
Suidae Dicoryphochoerus haydeni - 11.0 14.5 - -
Suidae Sivachoerus giganteus - 1.50 6.20 - -
Suidae Microstonyx major - 9.50 12.0 - - Suidae Sus pregrinus - 0.5 3.5 - - Suidae Sivahyus punjabiensis - 2.50 9.00 - - Suidae Hypohyus sp. - 1.50 8.10 - - Suidae Potamochoerus palaeindicus - 3.30 7.50 - - Tetraconodontinae Conohyus sindiensis 10.3 10.3 13.1 14.0 10 Tetraconodontinae Tetraconodon magnus 9.30 9.30 10.0 11.2 3 Suinae Propotamochoerus hysudricus 6.50 6.50 10.2 10.2 29 Suinae Hippopotamodon sivalense 7.1 7.20 10.2 10.2 21 Suinae HippopotamodonY450 indt sp. 10.2 10.2 11.3 11.4 05 Suinae Potamochoerus theobaldi - 0.5 3.5 Suinae Sus D013 species 2.20 3.30 6.40 6.80 03 Suinae “Small suinae Y311 species” 9.40 10.0 10.0 11.4 01 Suinae “Small suinae Y406 species” 2.20 7.40 8.90 9.20 03
Doliochoerinae Schizochoerus gandakasensis 8.10 8.70 10.1 11.2 06 Anthracotheriidae Microbunudon milaensis - 11.2 16.8 - -
Anthracotheriidae Anthracotherium punjabiensis - 10.3 18.3 - -
Anthracotheriidae Hemimeryx sp. 3.60 6.20 13.6 14.0 40 Anthracotheriidae Merycopotamus dissimilis 2.20 3.30 5.80 6.10 02 Anthracotheriidae Merycopotamus medioximus - 8.6 10.4 - - Anthracotheriidae Merycopotamus nanus - 11.3 13.9 - - 217
Appendix 9 (continued) Family/ Species Biostratigraphic Ranges N O SubFamily/Tribe ILA LO FO IFA I (Ma) (Ma) (Ma) (Ma) Hippopotamidae Hexaprotodon sivalensis 2.20 3.35 5.90 6.10 04
Tragulidae Dorcabune nagrii 8.40 8.50 10.4 11.2 10
Tragulidae Dorcabune anthracotheroides 10.3 10.5 12.8 14.0 09
Tragulidae Dorcatherium “Y270 sp.” 8.60 9.20 11.1 11.2 06
Tragulidae Dorcatherium nagrii 2.20 6.80 9.30 9.50 12
Tragulidae Dorcatherium “Y259 sp.” 10.3 10.4 12.3 14.0 03
Tragulidae Dorcatherium “Y311 species” 7.10 7.20 10.3 10.3 21
Tragulidae Dorcatherium “Y373 species” 6.50 6.80 7.80 7.80 05
Tragulidae Dorcatherium “Y457 species” 7.30 7.30 7.30 7.70 01
Tragulidae Dorcatherium cf. majus 10.5 10.7 11.3 11.4 03 Tragulidae Dorcatherium majus 6.90 7.00 10.4 10.6 19 Tragulidae Dorcatherium minus - 5.5 13.9 - - Tragulidae “Tragulidae/ L101 Indet sp.” 2.20 3.30 5.90 6.10 04 Tragulidae Archeotragulus - 16.0 18.0 - - Progiraffinae Progiraffa exigua 14.0 18.0 Paleotraginae Palaeotragus V55 - 11.0 18.0 - - Girrafinae Giraffa punjabiensis 2.20 7.20 8.90 9.10 03 Girrafinae Giraffa Priscilla - 11.0 15.0 - - Girrafinae Giraffa sivalensis - 1.50 6.5 - - Sivatheriinae Bramatherium megacephalum 7.10 7.10 10.3 10.3 26 Sivatheriinae Giraffokeryx punjabiensis 10.3 10.3 13.6 14.0 20 Sivatheriinae Hydaspith. Megacephalum - 5.20 10.2 - - Sivatheriinae Hydaspitherium magnum - 1.50 3.30 - - Sivatheriinae Sivatherium giganteum - 1.5 2.2 - - Sivatheriinae Helladotherium grandi - 3.50 10.3 - - 218
Appendix 9 (continued) Family/ Species Biostratigraphic Ranges N O SubFamily/Tribe I
ILA LO FO IFA (Ma) (Ma) (Ma) (Ma) Boselaphini Miotragocerus sp. - 4.50 10.2 - - Boselaphini Miotrag. salmontanus small sp. - 5.5 8.0 - - Boselaphini Miotragocerus gluten older - 8.0 13.7 - - Boselaphini Miotragocerus gluten younger - 5.0 8.0 - - Boselaphini Tragoporax browni older - 7.50 10.2 - - Boselaphini Tragoporax browni younger - 4.50 7.50 - - Boselaphini Pachyportax latidens - 3.50 10.3 - - Boselaphini Selenoportax vexillarius - 9.8 10.3 - - Boselaphini Protragocerus gluten - 10.8 13.9 - - Boselaphini Stresiportax sp. - 11.0 14.0 - - Boselaphini Helicoportax praecox - 10.8 14.2 - - Boselaphini Helicoportax tragelaphoides - 10.3 13.6 - - Boselaphini Helicoportax sp. - 12.5 13.5 - - Boselaphini Sivoreas eremite 10.3 10.7 11.3 14.0 02 Boselaphini Sivoreas gradiens - 10.3 12.0 - - Boselaphini Elachestoceros khaurestanensis 7.30 7.40 11.5 11.4 16 Boselaphini Selenoportax sp. 7.90 7.90 10.2 10.3 12 Boselaphini Selenoportax falconeri - 8.9 9.6 - - Boselaphini Selenoportax giganteus - 4.5 8.8 - - Boselaphini Tragoceridus sp. 2.20 6.20 11.2 11.2 22 Boselaphini Kubanotragus skolovi - 13.8 13.8 - -
Boselaphini cf. Eotragus “Y166 Indet sp.” 3.60 8.70 9.30 9.90 02 Boselaphini Eotragus noyie - 4.5 18.3 - Boselaphini Tragoportax salmontanus 7.30 8.40 9.30 9.90 03 Boselaphini Ruticeros pugio - 2.20 5.5 - - 219
Appendix 9 (continued) Family/ Species Biostratigraphic Ranges N O SubFamily/Tribe I
ILA LO FO IFA (Ma) (Ma) (Ma) (Ma) Boselaphini Paleohypsodontus zinensis - 13.2 20.0 - - Boselaphini Medium Boseliphini/Y581 Indet sp 6.30 7.00 7.00 7.10 01 Boselaphini 2-Keel-a large bovid species - 2.5 7.5 - - Boselaphini Medium Boseliphini/Y195 Indet sp 8.10 9.30 9.30 9.90 01 Boselaphini Large Boseliphini/Y927 Indet sp.’’ 7.30 7.30 8.00 8.10 02 Bovidae “Bovidae/Y166 Indet species” 8.40 8.70 8.70 9.10 01 Bovidae “Bovidae/Y905 Indet species” 7.40 7.80 7.80 7.80 01 Bovidae Small Bovidae/Y581 Indet sp. 6.30 7.00 7.00 7.10 01 Bovidae “Bovidae/Y545 Indet sp” 7.40 8.00 8.00 8.40 01 Tragilaphini Tragilaphini/D013species 5.80 6.20 6.40 6.70 03 Bovini Proamphibos lycrymans - 1.50 6.50 - - Bovini Proamphibos sp. - 1.5 9.9 - - Bovini Proamphibos kashmiricus - 0.50 2.6 - - Bovini Hemibos triquetricornis - 0.5 2.5 - - Bovini Hemibos antilopinus - 0.6 2.5 - - Bovini Hemibos acuticornis - 0.6 2.5 - - Bovini Babalus palaeindicus - 0.5 2.5 - - Bovini Bos acutifrons 0.6 2.5 - - Bovini Bos sp. - 0.50 3.50 - - Bovini Leptobos falconeri - 0.5 2.5 - - Bovini Bison sivalensis - 0.50 3.50 - - Bovini Bison crossicornis - 0.50 3.50 - - Reduncini “?Reduncini/D013 species” 2.20 6.40 6.40 6.90 01 Reduncini Sivacobus palaeindicus - 0.6 3.50 - - Reduncini Vishnucobus patulicornis - 0.6 2.5 - - 220
Appendix 9 (continued) Family/ Species Biostratigraphic Ranges N O SubFamily/Tribe I
ILA LO FO IFA (Ma) (Ma) (Ma) (Ma) Reduncini Hydaspicobus auritus - 0.5 5.5 - - Reduncini Indoredunca sterilis - 0.5 6.5 - - Reduncini Sivadenota sepulta - 0.50 3.50 - - Reduncini Kobus porrecticornis - 1.5 6.5 - 08 Hippotragini Hippotragus bohlini - 0.5 2.6 - - Hippotragini Hippotragus brevicornis - 0.5 2.6 - - Hippotragini Oryx sivalensis - 0.5 2.5 - - Hippotragini “?Hippotragini/Y453 species” 2.20 6.20 6.80 6.90 03 Caprini Sivacapra sivalensis 0.6 3.5 -
Caprini Caprotragoides potwaricus - 11.0 14.0 - - Antilopini Gazella sp. 2.20 6.20 11.3 14.0 23 Antilopini Gazella lydekkeri - 1.5 10.5 - - Antilopini cf. Prostrepsiceros vinayaki 7.30 7.40 7.60 7.90 04 Antilopini cf. Protragilaphus skouzesi 8.80 8.80 8.80 8.80 01 Antilopini Antilope subtorta - 1.50 3.50 - - Antilopini Antilope planicornis - 1.50 10.5 - - Antilopini Antilope cervicapra - 0.5 1.5 - - Antilopini Antilope intermedia sp.nov. - 0.5 2.5 - - Alcephini Demalops palaindicus - 0.5 2.2 - - Cervidae Rucervus simplicedense - 0.5 5.30 - - Cervidae Cervus triplidense - 1.5 5.20 - - Cervidae Cervus rewati - 1.50 5.20 - - Cervidae Cervus sivalensis - 0.6 5.50 - - Cervidae Cervus punjabiensis - 0.5 5.30 - - Hipparionini “Cormohipparion” sp. - 10.2 10.8 - - 221
Appendix 9 (continued) Family/ Species Biostratigraphic Ranges N O SubFamily/Tribe I
ILA LO FO IFA (Ma) (Ma) (Ma) (Ma) Hipparionini Sivalhippus nagriensis - 9.0 10.4 - - Hipparionini Sivalhippus theobaldi - 7.7 9.4 - - Hipparionini Sivalhippus perimense - 7.1 9.0 - - Hipparionini Hipparion sp. small 9.6 10 - - Hipparionini Sivalhippus anwari sp. nov 3.7 7.3 - - Hipparionini Cremohipparion antelopinum 3.7 8.9 - - Equidae Equus sivalensis - 1.50 3.50 - - Equidae Equus sp. - 1.50 3.50 - - Rhinocerotidae Brachypotherium perimense - 6.90 18.3 - - Rhinocerotidae Brachypotherium fatehjangense - 7.0 18.0 - - Rhinocerotidae Aceratherium blanfordi - 11.0 14.0 - - Rhinocerotidae Aceratherium sp. - 10.3 11.0 - - Rhinocerotidae Didermoceros aff. sumatrensis - 11.0 14.0 - - Rhinocerotidae Didermoceros aff. abeli - 11.2 14.0 - - Rhinocerotidae Chilotherium intermedium - 5.50 13.5 - Rhinocerotidae Caementodon oettingenae - 10.0 14.2 - - Rhinocerotidae Hispanotherium matritense - 10.3 14.0 - - Rhinocerotidae Gaindatherium vidali - 10.1 12.2 - - Rhinocerotidae Gaindatherium browni - 11.0 15.3 - - Rhinocerotidae Alicornops aff. laogouense - 12.0 16.0 - - Rhinocerotidae Alicornops complanatum - 3.10 10.0 - - Rhinocerotidae Rhinoceros sivalensis - 1.30 3.80 - - Rhinocerotidae Rhinoceros sondaicus - 1.0 3.20 - - Rhinocerotidae Punjabitherium cf. platyrhinus - 1.5 3.10 - - Chalicotherini Chalicotherium salinum 8.00 8.00 12.9 14.0 13
ILA= Inferred last appearance, LO= Last occurrence, FO= First occurrence, IFA=Inferred first appearance, NOI= Number of intervals in which observed. First and last occurrence data are from Sarwar (1974, 1977); Akhtar (1992); Ahmad (1995); Solounias et al. (1995); Barry et al. (2002); Lihoreau et al. (2004); Gaffar, (2005); Khan, (2006); Lihoreau et al. (2007); Badgley et al. (2008); Khan (2009); Khan et al. (2010); Gaffar et al. (2010, 2011); Khan and Akhtar (2011); Wolf et al. (2013); Khan and Akhtar (2014).
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Appendix 10; Mesowear database of extant species of ungulates Family Species Author Occlusal Cusp Shape (CS) C C (Abbrev) Relief o o (OR) d d L % H % S % R % B % 1 2
Cervidae Alces alces (AA) Linnaeus, 0.0 100 100 0.0 0.0 T B 1758 Bovidae Ammodorcas Thomas, 0.0 100 28.5 71.4 0.0 N B clarkei (EI) 1891 Antilocap- Antilopcaora Ord, 1815 4.0 96.0 88.6 11.3 0.0 N B ridae americana(AM) Bovidae Boocercus Ogilby, 0.0 100 44.4 55.5 0.0 N B euryceros (BE) 1837 Cervidae Capreolus Linnaeus, 4.0 96.0 72.0 25.0 2.9 N B capreolus (OL) 1758 Bovidae Cephalophus Gray, 1846 7.3 93.0 32.1 60.7 7.1 m B dorsalis (DR) Bovidae Cephalophus Smith, 0.0 100. 16.6 83.3 0.0 m B natalensis (NA) 1834 Bovidae Cephalophus niger Gray, 1846 9.0 91.0 35.4 61.2 3.2 m B (NI) Bovidae Cephalophus Gray, 1846 18. 82.0 25.0 70.4 4.5 m B nigrifron (NG) Bovidae Cephalophus Afzelius, 20. 80.0 0.0 94.8 5.1 m B silvicultor (SL) 1815 Procaviid- Dendrohyrax Smith, 0.0 100 50.0 50.0 0.0 m B ae arboreus (DA) 1834 Procaviid- Dendrohyrax Fraser , 18. 82.0 46.4 53.5 0.0 m B ae dorsalis (DD) 1855 Rhinocer- Dicerorhnus Fisher, 0.0 100. 80.0 20.0 0.0 T B otidae sumatrensis (DS) 1814 Rhinocer- Diceros Linnaeus, 0.0 100. 94.1 5.8 0.0 T B otidae bicornis(DB) 1758 Giraffidae Giraffa camelo- Linnaeus, 6.0 94.0 73.7 26.2 0.0 T B pardalis(GC) 1758 Procaviid- Heterohyrax Gray, 1846 64. 36.0 81.8 18.1 0.0 m B ae brucei(HB) Tragulida-e Hyaemoschus Ogilby, 0.0 100. 16.6 83.3 0.0 m B aquaticus(HY) 1837 Bovidae Litocarnius Brooke, 4.0 96.0 33.3 66.6 0.0 N B walleri(LW) 1841 Cervidae Odocoileus Rafinesqu, 0.0 100 72.7 27.2 0.0 T B hemionus (OH) 1817
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Appendix 10 (continued) Family Species Author Occlusal Cusp Shape (CS) C C o o (Abbrev) Relief (OR) d d 1 2 L % H % S % R % B % Cervidae Odocoileus Zimmerm 0.0 100 88.8 11.1 0.0 T B virginanus(OV) ann, 1780 Girrafidae Okapia Ssclater, 0.0 100 87.5 12.5 0.0 T B johnstoni(OJ) 1901 Rhinocer- Rhinoceros Desmarest 0.0 100 100. 0.0 0.0 T B otidae sondaicus(RS) , 1822 Bovidae Tragilaphus Pallas, 0 100 0.0 100. 0.0 N B strepsiceros (TS) 1766 Bovidae Alcelaphus Pallas, 43 57.0 3.2 66.6 28.0 T G buselaphus(ab) 1766 Bovidae Alcelaphus Lichtenst- 18 82.0 5.8 82.3 11.7 N G lichtensteinii(al) ein, 1812 Bovidae Bison bison(bb) Linnaeus 100 0.0 0.0 26.6 73.3 T G ,1758 Rhinocer- Ceratotherium Burchell, 100 0.0 0.0 72.0 28.0 T G otidae simum (cs) 1817 Bovidae Connochaetes Lichtenst- 45 55.5 15.3 55.7 28.8 T G taurinus (ct) ein, 1812 Bovidae Damaliscus lunatus Burchell, 80 20.0 20.0 60.0 20.0 T G (dl) 1817 Equidae Equus burchelli Gray, 100 0.0 27.0 39.3 33.6 T G (eb) 1846 Equidae Equus grevyi (eg) Oustalet, 100 0.0 34.4 41.3 24.1 T G 1882 Bovidae Hippotragus Desmarest 15 85.0 3.8 96.1 0.0 T G equines (he) , 1822 Bovidae Hippotragus Harris, 15.0 85.0 0.0 85.0 15.0 T G niger(hn) 1838 Bovidae Kobus Ogilby, 4.0 96.0 0.0 100 0.0 T G ellipsiprymnus(ke) 1837 Bovidae Redunca redunca Pallas, 9.0 91.0 6.4 90.9 2.5 T G (rr) 1767 Bovidae Aapyceros Lichtenste 0.0 100. 35.2 64.7 0.0 T M melampus(Me) in, 1812 0 Bovidae Antidorcas Zimmerm 4.0 96.0 73.0 26.9 0.0 N M marsupialis (Ma) ann, 1780 Cervidae Axis axis(Ax) Erxleben, 21.0 79.0 6.9 67.4 25.5 N M 1777 Cervidae Axis parcinus(Ap) Zimmerm 12.0 88.0 4.1 95.8 0.0 N M ann, 1780 224
Appendix 10 (continued) Family Species Author Occlusal Cusp Shape (CS) C C (Abbrev) Relief (OR) o o d d 1 2 L % H % S % R % B % Bovidae Boselaphus Pallas, 13.0 87.0 0.0 100. 0.0 N M tragocamelus (Btr) 1766 Bovidae Budorcas taxicolor Hadgson, 5.0 95.0 42.1 57.8 0.0 N M Bt 1850 Camilidae Camelus Linnaeus,1 0.0 100 31.2 68.7 0.0 N M dromedaries(Cd) 758 Bovidae Capra ibex(Ci) Linnaeus,1 3.0 97.0 54.1 37.5 8.3 N M 758 Bovidae Carpicornis Bechstein, 0.0 100 45.4 50.0 4.5 T M sumatraensis (Cs) 1799 Cervidae Cervus Linnaeus, 0.0 100 47.3 52.6 0.0 T M canadensis(Cc) 1758 Cervidae Cervus Curvier, 33.0 67.0 12.0 64.0 24.0 N M duvauceli(Cd) 1823 Cervidae Cervus unicolor Kerr, 1792 9.0 91.0 14.2 80.9 4.7 N M (Cu) Bovidae Gazelle granti(Gg) Brooke, 12.0 88.0 50.0 50.0 0.0 T M 1841 Bovidae Gazelle thomsoni Gunther, 12.0 88.0 55.4 43.1 1.3 T M (Gt) 1884 Camilidae Lama glama (Lg) Linnaeus,1 0.0 100. 28.1 68.7 3.1 N M 758 0 Camilidae Lama vicugna Molina, 0.0 100. 41.6 58.3 0.0 N M (Lv) 1782 0 Bovidae Ourebia ourebi Zimmerma 4.0 96.0 21.8 77.3 0.7 N M (Oo) nn, 1780 Bovidae Ovibos moschatus Zimmerma 19.0 81.0 57.6 42.3 0.0 T M (Om) nn, 1780 Bovidae Ovis Shaw,1804 13.0 87.0 48.3 51.6 0.0 N M canadensis (Oc) Procoviid- Procavia capensis Pallas, 54.0 46.0 62.5 37.5 0.0 m M ae (Pc) 1766 Bovidae Redunca Afzelius, 14.0 86.0 0.0 100. 0.0 N M fulvorufula (Rf) 1815 Rhinocer- Rhinoceros Linnaeus, 0.0 100. 80.0 20.0 0.0 N M otidae unicornis (Ru) 1758 Bovidae Saiga tatarica Linnaeus, 60.0 40.0 60.0 40.0 0.0 N M (St) 1758 Bovidae Syncerus caffer Sparman, 0.0 100. 0.0 93.5 6.4 N M (Sc) 1799 225
Appendix 10 (continued) Family Species Author Occlusal Cusp Shape (CS) C C (Abbrev) Relief (OR) o o d d 1 2 L % H % S % R % B % Bovidae Taurotragus oryx Pallas, 0.0 100. 50.0 50.0 0.0 T M (To) 1766 Bovidae Tetracerus Blainville, 9.0 91.0 28.5 71.4 0.0 N M quadricornis (Tq) 1816 Bovidae Tragelaphus Gray, 1849 0.0 100. 35.0 65.0 0.0 N M angasi(Ta) 0 Bovidae Tragelaphus Blyth, 1869 0.0 100. 61.2 38.7 0.0 N M imberbis (Ti) 0 Bovidae Tragelaphus Pallas, 0.0 100. 51.0 48.9 0.0 T M scriptus (Ts) 1766s 0
Frequency of mesowear variables in the recent comparison taxa (data from Fortelius and Solounias 2000), and recent Equus burchelli Gray, 1824 (ebN and ebS) investigated in this study. Abbreviations: low= percent low occlusal relief; high= percent high occlusal relief; sharp= percent sharp cusps; round= percent round cusps; blunt, percent blunt cusps; code 1: m=“mabra group” after Fortelius and Solounias (2000); n= neutral species; t= species with typical dietary adaptation; r= reference samples investigated; code 2: classification of dietary preferences according to the conservative classification by Fortelius and Solounias (2000); G= grazer; M= mixed feeder; B= browser.
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Appendix 11; Microwear database of extant species of ungulates
Taxon N Hyps A.P A.S % CS %LP % G %FS %CS %MS FRUIT-DOMINATD BROWSERS Cephalophus 07 Bra 33 9.43 100 71.43 57.14 42.86 0. 57.14 niger (CG) Tapirus 22 Bra 27.7 17.23 77.27 86.36 4.55 22.73 22.7 54.54 terrestris(TERR) Cephalophus 21 Bra 39 19.55 76.19 85.71 80.95 0 61.9 38.10 dorsalis (DR) Cephalophus 15 Bra 20 19.83 66.67 73.00 26.67 33.33 0.00 66.67 silvicator (SL) Okapiajohnstoni 25 Bra 34.6 20.68 84 84 56 8 56 36 (OJ) Tragulus sp. 11 Bra 30 22.09 54.55 100 0.00 9.09 63.6 27.27 (TRA) Moschusmos 3 Bra 37 25 100 100 33.33 0.00 100 0.00 chiferus (MO) Tapirus 20 Bra 27.85 25.05 75 90 10 25 35 40 bairdi (TBA) Cephalophus 4 Bra 32 27 0 100 0 0 25 75 natalensis(NA) LEAF BROWSERS Boocercus 15 Bra 6.77 5.13 13.33 53.33 0 100 0 0 euryceros(BE) Tragelaphus 19 Bra 16.84 7.08 22.22 27.78 47.37 83.33 0 16.67 imberbis (TI) Giraffa 28 Bra 5.00 8.66 54.17 20 0 91.67 0 8.33 camelopardalis( GC) Camelusbacteria 3 Hyp 79.83 9.17 100 66.67 66.67 33.33 33.3 33.33 nus(CB) 3 Alcesalces(AA) 9 Bra 27.15 10 44.44 0 0 66.67 0 33.33 Litocraniuswalle 22 Bra 27.14 11.8 9.09 27.27 45.46 90.91 0 9.09 ri (LW) Tragelaphusstre 11 Bra 20.09 12.6 18.18 63.64 36.36 90.91 9.09 9.09 psiceros (TT) Camelusdromed 7 Hyp 54.07 13.9 57.14 42.86 42.86 42.86 14.2 42.86 arius(CL) 9 Dicerosbicornis 11 Mes 8.5 14.3 95.24 0 0 85.71 9.52 4.76 (DB) Antilocapraamer 44 Hyp 25.21 15.5 53.49 79.55 69.77 37.21 0 62.79 icana(AM) 2 SEASONAL-REGIONAL MIXED FEEDERS Taurotragusoryx 20 Mes 39.8 12.23 55 55 42.11 44.4 16.6 38.89 (TO) 3 4 7 227
Appendix 11 (continued) Taxon N Hyps A.P A.S % CS %LP % G %FS %CS %MS Budorcastaxicolo 15 Hyp 7.97 13.73 33.33 56.25 0 100 0 0 r (BT) Gazellagranti 41 Hyp 20.5 14.77 87.81 53.66 39.02 39.0 9.76 51.22 (GG) Tragelaphusscript 31 Bra 19.0 15.87 19.36 19.36 22.58 70.9 16.1 12.90 us (TS) Lama vicugna 7 Hyp 30.4 16.21 28.57 57.14 85.71 28.6 42.9 28.57 (VI) Oviscanadensis(O 19 Hyp 13.2 16.71 31.58 42.11 47.37 15.9 15.9 68.42 C) Lama glama(LG) 6 Hyp 4.92 18.42 100 0 16.67 100 0 0 Gazellathomsoni 23 Hyp 24.0 18.57 80.95 57.14 30.44 14.3 14.3 71.43 (GT) Capricornis 11 Hyp 40.1 21.73 55.56 44.44 80 77.8 0 22.22 sumatraensis(CA) Boselaphus 9 Mes 23.6 25.72 55.56 44.44 88.89 44.4 22.2 33.33 tragocamelus(TR) Axis axis (AX) 43 Mes 13.6 28.28 75.61 9.76 9.76 52.5 22.5 25 Muntiacus 10 Bra 20.5 18.95 80 72.73 30 20 0 80 muntijak (MM) MEAL BY MEAL MIXEDFEEDERS Cervus unicolor 11 Mes 21.3 26.27 100 36.36 100 18.18 45.5 36.36 (CU) Ovibos 20 Hyp 28.5 29.68 15 100 85 50 0 50 moschatus (OM) Cervuscanad- 28 Bra 18.6 30.63 96.43 42.86 21.43 82.14 0 17.86 ensis (CC) Capra ibex (CI) 09 Hyp 7.56 25.2 30 50 20 10 0 90 GRAZERS Connochaetes 27 Hyp 4.93 20.6 44.44 40.74 7.41 44.44 7.41 48.15 taurinus (CT) Equus 51 Hyp 11.5 21.7 60 48.98 50 40 10 50 burchelli (EB) 2 Hippotragus 5 Hyp 5.90 22.7 0 60 0 20 40 40 niger (HN) Bison bison (BB) 18 Hyp 3.53 24.8 94.44 38.89 5.26 22.22 50 27.78 Tetracerusquad- 8 Mes 22.2 25.7 50 87.50 75.00 87.50 0 12.50 ricornins(TQ) Equus grevyi(EG) 11 Hyp 7.86 26.0 66.67 63.64 58.33 0 66.7 33.33 Alcelaphus 6 Hyp 13.6 29.6 12.50 75.00 62.50 0 12.5 87.50 buselaphus (AB) Cervus 26 Mes 15.8 30.50 60.00 16.00 0.00 12.00 4.00 84.00 duvauceli(CD) 228
Appendix 11 (continued) Taxon N Hyps A.P A.S % CS %LP % G %FS %CS %MS SUIDS Hylochoerus 3 - 23.3 17 100 0.0 0.0 100 0.0 0.0 mainertzhageni (HM) Potamochoerus 14 - 34.8 20.1 100 92.86 35.71 7.14 42.9 50 porcus (PP) Babyrousa 8 - 26.4 25.9 100 100 50 0 37.5 62.5 babyrousa (BB) ELEPHANTS Loxadonta 09 Hyp 15.9 24.50 100 100 100 0 100 0.0 africana (LA) Elephus maximus 7 Hyp 9.86 28.18 85.7 71.43 85.71 0 71.4 28.57 (EM)
Key to Microwear Symbols (headings) N= Number of individuals; Hyps = hypsodonty; Bra= brachydont; Mes= mesodont; Hyp= hypsodont; AP= Average Pits; AS= Average Scratches; % CS= percentage of coarse scratches; % LPs= percentage of large pits; % G= percentage of gouges; % FS= percentage of fine scratches; % CS= percentage of coarse scratches; %MS= percentage of mixed scratches.
229
Appendix 12; Morphometric and ecomorphic database of living species of ungulates. Taxon No. of obs. Total Tooth Volume Height Measures B.W Di et Area Ht. Mean Range H.I M3 M3 Ht Vol ARTIODACTYLA Antilocapridae M 50 6.52 3.64 4.61 19.23-13.22 16.04 ٭Antilocapra americana 11 1 Bovidae-Alcelaphini Aepyceros melampus 09 02 14.99 12.54-18.31 4.89 3.52 5.95 53 M Alcephalus buselaphus 16 01 32.17 26.94-41.02 5.23 4.60 10.58 136 G G 136 14.21 4.85 4.75 45.56-34.05 38.98 ٭Connochaetes gnou 5.0 1 Connochaetes taurinus 18 01 49.08 35.90-64.33 4.94 5.63 20.72 216 G Damaliscus dorcas 10 01 21.59 14.95-27.73 4.76 3.90 7.22 69 G Damaliscus hunteri 07 01 26.17 22.78-29.30 4.14 4.02 10.81 88 M Damaliscus lunatus 10 01 37.55 27.17-38.60 5.10 4.60 15.59 150 G Boselaphini Boselaphus 09 01 35.37 23.92-43.83 3.03 3.70 13.99 210 W tragocamelus Tetracerus quadricornis 07 02 3.77 2.66-4.93 2.89 2.89 1.50 17 W Bovini Bison bison 09 01 120.56 95.31-138.8 4.87 7.30 46.65 675 G G 625 32.54 7.28 6.12 0 91.01 ٭1 ٭Bison bonasus 1 Bos gaurus 06 01 101.98 87.06-111.6 3.69 6.20 41.04 755 M Bos indicus 02 01 73.05 57.71-88.28 4.43 5.71 27.29 600 M Syncerus caffer 10 01 77.56 5.50-99.44 3.00 4.77 29.48 620 M Caprini Ammotragus lervia 05 01 23.39 20.36-26.62 4.45 3.88 9.23 86 M Capra ibex 07 02 18.33 12.52-23.09 4.71 4.71 4.00 87 M Hemitragus jehmlahicus 05 01 18.96 13.80-23.21 4.95 4.11 8.22 91 M Ovis canadensis nelson 04 01 21.68 19.30-24.89 4.11 3.82 9.40 59 M Pseudois nayaur 01 01 14.34 13.68-15.98 5.30 4.08 6.73 59 M Cephalophini Cephalophus dorsalis 04 01 4.55 4.18-5.02 1.15 1.30 1.44 20 S Cephalophus monticola 08 01 0.91 0.72-1.15 1.90 0.80 0.30 5.5 S Cephalophus silviculator 09 01 14.93 11.82-17.7 2.23 2.32 5.41 61 B Cephalophus spadix 04 01 5.38 4.90-5.94 1.95 1.48 1.88 57 S Silvicapra grimmia 02 02 4.42 4.11-4.79 2.97 1.72 1.46 13 B Gazellini Ammodorcas clarkei 03 02 2.64 2.29-2.97 2.23 1.25 1.08 28 H Antilope cervicapra 04 01 11.53 8.39-13.49 5.14 3.65 5.07 37 M Antidorcas marsupialis 11 01 10.33 8.83-13.54 4.89 3.18 4.45 31 M Gazella dorcas 07 03 4.05 2.77-5.24 3.62 1.88 1.56 23 M 230
Appendix 12 (continued) Taxon No. of Total Tooth Volume Height Measures B.W Di obs. et Area H Mean Range H.I M3 M3 t. Ht Vol Gazella granti 07 02 11.69 10.02-15.09 3.45 2.83 5.55 62 M Gazella thomsoni 21 01 9.28 4.75-8.43 3.77 2.26 2.46 20 M Litocranius walleri (X) 15 03 1.79 1.35-2.27 1.32 0.78 0.66 43 H Procapra gutterosa 06 02 9.90 6.14-12.33 3.90 2.89 4.74 18 M Hippotragini Addax nasomaculatis 04 01 28.48 24.59-34.93 4.09 4.17 13.80 111 M Hippotragus equines 09 02 36.32 38.07-57.73 4.28 4.41 12.97 270 G Hippotragus niger 10 02 35.81 29.79-45.16 3.77 3.87 11.38 225 G Oryx gazelle 06 02 45.63 37.36-53.56 3.37 4.08 16.61 170 M Neotragini Dorcatragus megalatus 03 01 2.01 1.86-2.17 2.93 1.20 0.64 09 M Madoqua kirki 03 01 1.19 1.04-1.33 2.63 1.00 0.33 4.50 S Neotragus pygmaeus 06 02 0.79 0.62-0.96 2.94 1.06 0.38 3.50 S Neotragus moschatus 07 01 1.18 1.01-1.47 3.51 1.30 0.48 4.50 S Orebia orebi 10 02 4.26 3.42-5.22 3.80 2.05 1.72 18 M Oreotragus oreotragus 04 02 2.98 2.30-3.69 3.82 1.68 1.03 13.5 M Ramphicerus campestris 16 02 1.95 1.11-2.47 3.44 1.48 0.78 13.5 M Ramphicerus melanotis 08 02 1.73 1.27-2.15 2.64 1.11 0.59 10 M Reduncini Kobusellipsi prymnus 11 01 30.92 22.70-39.38 3.47 3.82 11.92 205 F Kobus kob 02 01 11.60 10.33-13.20 3.72 2.75 3.39 57.8 F Kobus lechwe 04 01 13.95 11.73-16.70 3.63 2.90 5.10 87 F Kobus vardoni 04 02 12.93 11.18-15.03 4.05 3.04 5.05 87 F Pelea capreolus 05 01 6.81 5.27-8.00 4.01 2.67 3.34 32 M Redunca arudinum 09 03 9.96 7.55-13.72 3.59 2.69 4.17 62 F Redunca julvorufula 14 02 7.97 5.75-13.84 3.79 2.77 3.38 31 M Rupicarini Budorcas taxicolor 06 01 47.05 39.33-55.40 3.43 4.43 19.23 250 M Capricornis sumatrensis 07 02 21.69 15.57-24.91 3.93 3.73 9.14 102 M Nemorhaedus goral 07 02 10.76 9.01-12.30 4.03 2.90 3.83 27 M Oreamus americanus 08 01 11.25 9.52-13.19 2.76 2.35 4.98 97 M M 364 23.04 4.50 3.69 0 61.42 ٭1 ٭Ovibos moschatus 1 M 50 4.28 3.10 4.56 9.02-8.28 8.63 ٭1 ٭Pantholops hodgsoni 1 Rupicapra rupicapra 05 02 6.22 5.31-7.47 4.19 2.47 2.37 39 M Saiga tatarica 05 02 10.70 9.74-11.73 5.29 3.28 4.69 45 M Tragilaphini Taurotragus oryx 10 01 63.84 41.72-95.97 2.91 4.43 27.64 511 M Tragelaphus angasi 12 01 12.26 9.34-15.61 2.52 2.34 5.36 91 W Tragelaphus buxtoni 02 01 22.26 21.24-23.22 1.95 2.48 8.88 183 W Tragelaphus euryceros 06 01 28.68 25.04-31.75 1.92 2.49 11.03 205 B 231
Appendix 12 (continued) Taxon No. of Total Tooth Volume Height Measures B.W Di obs. et Area Ht Mean Range H.I M3 M3 . Ht Vol Tragelaphus imberbis 07 01 12.24 9.44-14.46 1.97 1.91 4.99 77 W Tragelaphus scriptus 12 01 5.06 3.75-6.16 2.54 1.73 2.21 58 W Tragelaphus spekei 04 02 9.53 8.47-10.36 2.90 2.29 3.73 74 F Tragelaphus 12 01 35.55 23.67-43.83 2.29 3.07 13.60 215 B strepsiceros Camilidae Camelus bacterianus 03 02 133.3 116.17-157.43 2.98 5.87 62.93 550 M Camelus dromedaries 04 01 125.7 103.22-145.80 2.52 5.25 58.5 550 M Lama guanicoe 04 01 20.80 17.37-24.36 3.46 3.42 8.89 110 M Vicugna vicugna 09 02 13.50 8.73-12.95 4.33 3.55 6.67 50 M Cervidae Alces alces (X) 07 03 66.55 53.91-80.35 1.34 2.48 18.75 384 H Axis porcinus 04 01 7.38 7.12-7.77 2.53 1.85 2.94 42 M Blastocerus dichotomus 08 02 10.6 7.88-12.05 1.49 1.52 3.39 130 W (X) Capreolus capreolus 05 01 6.96 6.26-7.62 1.49 1.56 2.47 30 W Cervus canadensis(X) 09 01 39.69 31.04-48.25 1.96 2.76 13.33 325 W Cervus elephus scottius 08 01 17.01 12.77-20.21 2.11 2.17 5.75 153 W Cervus nippon 07 01 9.06 7.82-10.81 2.79 2.18 3.29 53 W Cervus unicolor equinus 02 01 29.68 24.98-34.36 2.20 2.75 10.45 188 W Dama dama 04 02 9.80 7.43-11.72 2.01 1.85 3.65 55 W Elephus cephalophus 06 02 3.85 3.25-4.48 1.69 1.23 1.39 18 W Elaphurus davidianus 11 01 36.78 30.81-45.40 2.35 2.84 10.65 17.5 F Hippocamelus bisculus 06 04 8.63 6.19-10.43 1.94 1.61 2.85 50 W Hydrpotes inermis (X) 07 02 2.56 2.26-2.94 1.84 1.05 0.79 11 W Mazama mazama 05 02 3.23 2.43-3.79 1.30 0.90 0.92 20 B americana (X) Muntiacus muntjak 06 02 4.66 3.58-5.82 1.81 1.34 1.61 25 W vaginalis Muntiacus reevesi 02 02 2.55 2.22-2.78 1.12 0.84 0.91 13 W Odocoileus hemionus(X) 07 03 9.66 8.18-11.51 1.59 1.17 2.96 74 B Odocoileus virginianus 16 03 6.43 5.05-8.67 1.23 1.17 2.01 52 B (X) Ozotoceros bezoarticus 03 03 7.02 5.20-8.78 2.12 1.61 2.50 37 M Pudu mephistopheles 03 01 1.53 1.42-1.61 1.61 0.90 0.52 7 B Pudu pudu 05 02 1.96 1.57-2.69 1.62 1.00 0.66 9 B Rangifer tarandus (X) 05 01 12.33 10.66-14.29 1.52 1.40 2.70 145 B Giraffidae Giraffa camelopardalis 25 02 60.91 43.63-75.66 1.20 2.26 16.61 1075 H (X) 232
Appendix 12 (continued) Taxon No. of obs. Total Tooth Volume Height Measures B.W Di et Area Ht. Mean Range H.I M3 M3 Ht Vol Okapia johnstoni (X) 05 04 33.86 28.52- 1.18 1.96 10.70 250 H 39.18 Hippopotamidae Choeropsis liberiensis 02 01 35.05 25.33- 1.37 2.28 12.93 240 B 44.62 Hippopotamus 05 01 367.72 308.87- 1.91 6.38 146.4 3200 F amphibious 422.14 8 Suidae Babyrussa babyrussa (X) 07 02 6.37 4.89-8.35 1.03 1.11 2.70 85 O Hylochoerus 06 04 21.44 15.63-31.41 1.36 2.10 12.24 235 B meinerizhageni(X) Phacochoerus 04 01 31.66 21.33-38.02 3.99 4.96 24.25 100 G aethiopicus Potamochoerus 05 01 12.85 11.35-14.24 1.34 1.62 5.41 103 O porcus (X) Susscrofa 06 01 16.22 12.30-20.01 1.28 1.70 9.00 120 O cristatus (X) Tayassuidae O 38 4.84 1.40 1.06 0 14.53 ٭1 ٭Catagonus wagneri (X) 1 Tayas supecari (X) 04 01 8.44 7.49-9.52 1.23 1.23 2.92 32 O Tayas sutajacu(X) 04 01 5.09 4.62-5.42 1.00 0.97 1.70 22 O Tragulidae Hyaemoschus 06 02 1.96 1.36-2.13 1.30 0.74 0.56 12.5 S aquaticus(X) Tragulus javanicus (X) 05 05 0.62 0.34-0.64 1.47 0.53 0.18 2.5 S Tragulus memmina (X) 04 04 0.77 0.60-0.96 1.72 0.67 0.28 ? S PERISSODACTYLA Equidae Equus aesinus 04 02 115.48 99.27- 8.73 6.90 15.95 220 G 133.55 Equus burchelli 23 01 111.92 86.43- 5.83 6.76 21.02 235 G 155.76 Equus grevyi 06 01 171.86 142.77- 5.80 7.90 32.23 400 G 197.77 Equus hemionus 05 01 159.02 141.43- 5.79 7.47 25.92 290 G 177.42 Equus kiang 05 01 172.37 149.79- 6.08 7.90 28.36 300 G 192.37 Equus przewalski 02 01 207.18 200.48- 5.70 7.64 30.87 350 G 213.79 233
Appendix 12 (continued) Taxon No. of obs. Total Tooth Volume Height Measures B.W Di et Area Ht. Mean Range H.I M3 M3 Ht Vol Equus zebra 09 02 171.88 133.67- 7.06 8.75 30.80 260 G 213.42 Rhinocerotidae Ceratotherium simum 06 02 535.21 404.12- 3.90 10.14 151.0 3000 G 758.39 9 Dicerorhinus sumatrensis 02 01 149.42 142.15- 1.67 3.27 26.88 800 B 157.93 Diceros bicornis 08 02 250.38 203.23- 2.24 5.05 54.79 1800 B 312.97 Rhinoceros sondaicus 02 02 193.21 157.05- 1.72 4.20 43.60 1400 B 220.42 Rhinoceros unicornis 04 03 226.92 179.73- 1.59 4.12 56.11 2220 W 276.86 Tapiriidae Tapirus indicus (X) 02 01 41.81 38.30-41.86 0.76 1.60 9.65 275 H Tapirus terrestris (X) 05 02 28.84 25.05-34.10 0.92 1.64 7.220 240 H HYRACOIDEA Dendrohyrax dorsalis(X) 08 02 0.55 0.41-0.78 1.53 0.55 0.12 4.50 H Heterohyrax johnstoni(X) 08 02 0.37 0.31-0.42 1.52 0.47 0.09 3 M Procavia capensis 04 02 0.67 0.42-0.91 1.69 0.61 0.17 4 G PROBOSCIDEA Loxodonta africana - - 7610 - 2.50 20.0 4160 6000 M
Key to dietary symbols G= dry grass grazer; F= fresh grass grazer; M= mixed feeder in open habitat; W= mixed feeder in closed habitat; B= unspecialized browser; S= selective browser; H= high level browser; O= omnivore (X) after species name indicates that dental height values were available by direct measurements (i.e., no X-ray was taken) (Source; Janis, 1988). by the number of observations means that only one individual specimen was used in making all (٭) A star the measurements. Data on Loxodonta africana compiled from Laws (1966) by Dr. Louise Roth.