TAXONOMIC STUDIES OF FOSSIL EVEN AND ODD- TOED MAMMALS FROM THE MIOCENE ROCKS OF DHOK BUN AMEER KHATOON, DISTRICT CHAKWAL, PUNJAB, PAKISTAN

By KHIZAR SAMIULLAH

A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF Doctor of Philosophy In the Faculty of the Life Sciences, University of the Punjab, Lahore, Pakistan

Supervisor PROF. DR. MUHAMMAD AKHTAR

Department of Zoology University of the Punjab, Quaid-e-Azam Campus Lahore, Pakistan.

(2010)

IN THE NAME OF ALLAH, MOST GRACIOUS, MOST MERCIFUL

DEDICATION

Dedicated to MY PARENTS BROTHERS, SISTERS AND MY WIFE i

ABSTRACT

Fossil site Dhok Bun Ameer Khatoon (32o 47' 26.4" N, 72° 55' 35.7" E) yielded a significant amount of mammalian assemblage including four families of even-toed fossil mammal (Suidae, Tragulidae, Giraffidae, and Bovidae) and one family of odd-toed (Rhinocerotidae) of the Late Miocene. This newly discovered site has well exposed Chinji and Nagri formation and has dated approximately 14.2 Ma-9.5 Ma. This age agrees with the divergence of different mammalian genera and is important Palaeoecologically, Palaeogeographically and Palaeoclimatologically. Sedimentological evidence of the site supports that this is deposited in locustrine or fluvial environment, as Chinji formation is composed primarily of mud-stone while the Nagri formation is sand dominated. Palaeoenvironmental data indicates that Miocene climate of Pakistan was probably be monsoonal as there is now a days. Mostly the genera recovered from this site resemble with the overlying younger Dhok Pathan formation of the Siwaliks while the size variation in dentition is taxonomically important for vertebrate evolutionary point of view and this is the main reason to conduct this study at this specific site to add additional information in the field of Palaeontology. A detailed study of fossils even and odd-toed mammals found in Miocene rocks exposed at Dhok Bun Ameer Khatoon was carried out. Over all one hundred and twenty specimens were collected during field trips from which forty two specimens are well preserved and identified and are described in this thesis. Two specimens belonging to Gaindatherium browni, five specimens belonging to Listriodon pentapotamiae, four specimens belonging to Dorcatherium majus, one specimen belonging to Dorcatherium cf minus, two specimens belonging to Giraffa priscilla, twenty one specimens belonging to Giraffokeryx punjabiensis, three specimens belonging to Gazella sp. and three specimens belonging to Eotragus sp. Dhok Bun Ameer Khatoon is the new locality which is discovered in detail first time in Pakistan by the present author. The collection comprises isolated upper and lower teeth and fragments of maxillae and mandibular ramii.

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ACKNOWLEDGEMENTS

All praises to omnipotent Allah, who bestowed me the wisdom and courage to complete this palaeontological research work successfully. And Darood-o-salam to Hazrat Muhammad (Peace Be Upon Him). Who is forever a path of guidance for mankind and enables us to recognize our creator and to understand the philosophy of life.

It is my privilege and honour to express the deepest gratitude to my dignified supervisor, Prof. Dr. Muhammad Akhtar for his inspiring guidance, unprecedented help and intellectual suggestions. Without his help it would not have been possible for me to produce this work. Thanks and gratitude to my Father, Mother, Brothers and Sisters, as without their prayers and support I was unable to accomplish this work. Dr. Abdul Ghaffar and Dr. Muhammad Akbar Khan are thanked for their regular visits with me as well as for their precious advices during field work.

Mr. Saeed-ur-Rehman Deputy Director Fisheries is thanked for urging me for Ph. D. and Mr. Ghulam Murtaza, Mr. Muhammad Abid, Muhammad Hassan Siddiqi and Shaukat Bhatti are thanked for being patient with me. I would like to thank Abdul Majid Khan, Mr. Abdul Rehman and Mr. Muhammad Bilal for their support and help in photography.

The author is grateful to Engineer Jahangeer Siddique Bhatti, Sub Divisional Officer and Assad Shabbir, Lecturer institute of Mycology and Plant Pathology, for their helpful discussion and suggestions for the improvement of the manuscript. Hafiz Muhammad Nazir, my supper senior is also being thanked for his help to prepare my Ph. D. synopsis in my early days. I would like to thank Mr. Mahboob Iqbal, Assistant Professor for his kind co-operation during my research work. I offer my profound thanks to Dr. Muhamad Shafiq Ahmad for his absorbing attitude, useful intellectual suggestions and helping me in all the difficulties. I pay my cordial thanks to my loving and caring Brothers Humayoun Malik, Jawad Malik, Babar Malik, and Aamir Malik and sisters who prayed and encouraged me a lot. I will also ever remember the exceptional support and prayers of my wife, who patiently waited for the completion of this work.

I am also thankful to Mr. Maskeen Ali laboratory attendant for his help and continuous visits to study area with me. I am grateful to Mr. Maqsood Hussain (Lab. Attendant) for his assistance in the hours of need. iii

I would like to thank all the people from the village Dhok Bun Ameer Khatoon for their hospitality. Special thanks to Mr. Iftikhar and his son Mohsin for their help in my every field trip for fossil collections. If I forget to mention any person then I do apologise and say so lot of thanks whom I missed.

Khizar Samiullah Malik

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ABBREVIATIONS

Ma Million years ago MN European Mammal Neogene zone scale No. Number mm millimeter AMNH American museum of Natural History, New York, U.S.A. GSI Geological survey of India (Kolkata) GSP Geological survey of Pakistan (Islamabad) PUPC 08/11 Punjab University Palentological Collection, housed in the department of Zoology, Punjab University, Lahore, Pakistan (Institutional abbreviation). In the text the lower figure of the serial numbers denotes the collected number of the specimen of the respective years. I1 Upper Incisor 1 M 1 First upper or lower molar 2 M 2 Second upper or lower molar 3 M 3 Third upper or lower molar 2 P 2 Second upper or lower premolar 3 P 3 Third upper or lower premolar 4 P 4 Fourth upper or lower premolar L maximum preserved length W maximum preserved width H maximum preserved height W/L Width/Length ratio v

CONTENTS Abstract ...... i Acknowledgements ...... ii Abbreviations ...... iv Contents ...... v

List of Tables ...... viii List of Figures ...... ix List of plots ...... xii

CHAPTER 1 Introduction ...... 1 Geography of the Siwaliks ...... 1 Geology of the Siwaliks ...... 3 Lithology of the Siwaliks ...... 4 Sedimentology of the Siwaliks ...... 8 Taphonomy of the Siwaliks ...... 9 Siwalik Stratigraphy...... 11 Siwalik Chronostratigraphy ...... 14 Siwalik Biostratigrapy and Paleoenvironments ...... 16 Studied Section ...... 19 Methodology ...... 21 Tooth Morphology ...... 22 Objectives of Present Study ...... 24 Thesis Layout ...... 24

CHAPTER 2 Literature Review...... 25 Rhinocerotids ...... 26 Suids ...... 27 Tragulids ...... 28 Giraffids ...... 29 Bovids ...... 30 vi

CHAPTER 3 Systematic Palaeontology ...... 33 Genus: Gaindatherium Colbert, 1934 ...... 33 Gaindatherium browni Colbert, 1934 ...... 34 Description ...... 34 Discussion ...... 41 Genus: Listriodon Von Meyer, 1846 ...... 43 Listriodon pentapotamiae Falconer, 1868 ...... 44 Description ...... 44 Discussion ...... 59 Genus: Dorcatherium Kaup, 1833 ...... 61 Dorcatherium majus Lydekker, 1876 ...... 62 Description ...... 63 Discussion ...... 73 Dorcatherium cf. minus Lydekker, 1876 ...... 76 Description ...... 76 Discussion ...... 80 Genus: Giraffa Brisson, 1762 ...... 82 Giraffa priscilla Mathew, 1929 ...... 83 Description ...... 83 Discussion ...... 90 Genus Giraffokeryx Pilgrim, 1910 ...... 91 Giraffokeryx punjabiensis Pilgrim, 1910 ...... 91 Description ...... 92 Discussion ...... 127 Genus: Gazella Blainville, 1816 ...... 129 Gazella sp...... 130 Description ...... 131 Discussion ...... 139 Genus: Eotragus Pilgrim, 1939...... 141 Eotragus sp...... 142 Description ...... 142 Discussion ...... 152 vii

CHAPTER 4 Discussion of the Even and Odd-Toed Mammals ...... 154 Age and Correlation ...... 157 Conclusion ...... 163 REFERENCES ...... 164 APPENDIX ...... 191

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LIST OF TABLES

Table Page Title No. No. Table 1: Biostratigraphic interval-zones scheme with characteristics fauna of the Siwalik Group (and Murree Formation) of the 16 Potwar Plateau (Shah et al., 1985). Table 2: Comparative Measurements (in mm) of teeth referred to G. 38 browni Colbert, 1934. Table 3: Measurements (in mm) of teeth referred to L. pentapotamiae 53 Falconer, 1868. Table 4: Dental measurements (in mm) of different species of the genus 56 Listriodon. Table 5: Comparative Measurements (in mm) of molar teeth referred to 70 D. majus Lydekker, 1876. Table 6: Comparative Measurements (in mm) of left M1 referred to D. 79 cf. minus Lydekker, 1876. Table 7: Measurements (in mm) of molar teeth referred to Giraffa 88 priscilla Mathew, 1929. Table 8: Comparative measurements (in mm) of the cheek teeth 119 referred to G. punjabiensis Pilgrim, 1910. Table 9: Comparative measurements (in mm) of the cheek teeth of 136 Gazella sp. Table 10: Comparative measurements (in mm) of the cheek teeth of 147 Eotragus sp. Table 11: Dental measurements (in mm) of different species of the genus 147 Eotragus.

ix

LIST OF FIGURES

Figure No. Title Page No.

Figure 1 The Siwalik sediments along the foothills of Himalayas 2 (Chauhan and Gill, 2002). Figure 2 Reconstruction of Late Miocene Siwalik fluvial system 7 in plan view. Figure 3 Map of the Potwar Plateau (northern Pakistan) and a generalized stratigraphic section of the major Siwalik 13 formations showing succession and ages (source Behrensmeyer and Barry, 2005). Figure 4 Map of the studied section Dhok Bun Ameer Khatoon, 19 district Chakwal, Punjab, Pakistan. Figure 5 A-D Terminology of tooth crown elements in even and odd 23 toed mammals. Figure 6 G. browni, PUPC 08/14, an isolated lower right second 36 premolar. Figure 7 G. browni, PUPC 08/85, a right mandibular ramus with 37 M2-3. Figure 8 L. pentapotamiae, PUPC 08/9, right mandibular 47 fragment having P4-M3. Figure 9: L. pentapotamiae, PUPC 08/9, left mandibular 48 fragment having M1-3. Figure 10 L. pentapotamiae, PUPC 08/93, left third lower 49 premolar. Figure 11 L. pentapotamiae, PUPC 08/21, right fourth lower 50 premolar. Figure 12 L. pentapotamiae, PUPC 08/70, left first lower molar. 51 Figure 13 L. pentapotamiae, PUPC 08/24, right upper canine. 51 Figure 14 L. pentapotamiae, PUPC 08/9, left upper incisor. 52 Figure 15 D. majus, PUPC 08/23, right maxillary fragment having 66 M2-3. x

Figure No. Title Page No.

Figure 16 D. majus, PUPC 08/36, an isolated right second upper 67 molar. Figure 17 D. majus, PUPC 08/42, an isolated right second upper 68 molar. Figure 18 D. majus, PUPC 08/39, right first lower molar. 69 Figure 19 D. cf. minus, PUPC 08/90, left first upper molar. 78 Figure 20 Giraffa priscilla, PUPC 08/29, right second lower 86 molar. Figure 21 Giraffa priscilla, PUPC 08/10, left mandibular ramus 87 having M1-2. Figure 22 G. punjabiensis, PUPC 08/35, isolated left second 98 upper premolar.

Figure 23 G. punjabiensis, PUPC 08/97, isolated right second 99 upper premolar. Figure 24: G. punjabiensis, PUPC 08/33, isolated right third upper 100 premolar. Figure 25: G. punjabiensis, PUPC 08/34, isolated left third upper 101 premolar. Figure 26: G. punjabiensis, PUPC 08/43, isolated left third upper 102 premolar. Figure 27: G. punjabiensis, PUPC 08/98, isolated left third upper 103 premolar. Figure 28: G. punjabiensis, PUPC 08/26, isolated left fourth upper 104 premolar. Figure 29: G. punjabiensis, PUPC 08/96, isolated right fourth 105 upper premolar. Figure 30: G. punjabiensis, PUPC 08/31, isolated right first upper 106 molar. Figure 31: G. punjabiensis, PUPC 08/105, isolated right first upper 107 molar. Figure 32: G. punjabiensis, PUPC 08/113, isolated left first upper 108 molar. xi

Figure No. Title Page No.

Figure 33: G. punjabiensis, PUPC 08/99, isolated left second 109 upper molar. Figure 34: G. punjabiensis, PUPC 08/18, isolated right second 110 upper molar. Figure 35: G. punjabiensis, PUPC 08/28, isolated right second 111 upper molar. Figure 36: G. punjabiensis, PUPC 08/117, isolated right third 112 upper molar. Figure 37: G. punjabiensis, PUPC 08/38, isolated right third lower 113 premolar. Figure 38: G. punjabiensis, PUPC 08/94, isolated left third lower 114 premolar. Figure 39: G. punjabiensis, PUPC 08/95, isolated left third lower 115 premolar. Figure 40: G. punjabiensis, PUPC 08/45, isolated left first lower 116 molar. Figure 41: G. punjabiensis, PUPC 08/13, isolated right second 117 lower molar. Figure 42: G. punjabiensis, PUPC 08/11, isolated right third lower 118 molar. Figure 43: Gazella sp., PUPC 08/16, a right mandibular ramus 133 having M1-2. Figure 44: Gazella sp., PUPC 08/116, a left mandibular ramus 134 having M1-2. Figure 45: Gazella sp. PUPC 08/111, a left second upper molar. 135 Figure 46: Eotragus sp., PUPC 08/100, right mandibular ramus 144 having P3-M3. Figure 47: Eotragus sp., PUPC 08/101, left mandibular fragment 145 with M1-3.

Figure 48: Eotragus sp., PUPC 08/114, a right first lower molar. 146

xii

LIST OF PLOTS

Page Plot No. Title No. Plot 1 Dhok Bun Ameer Khatoon even and odd-toed families and 20 number of species described in this thesis. Plot 2 Dhok Bun Ameer Khatoon even and odd-toed species and 20 number of specimens described in this thesis. Plot 3a-c Bivariate scatter graphs showing comparison of the studied 39 specimens with the type specimens. Plot 4 Width/length indexes of the cheek teeth of G. browni. 40 Plot 5 Size variation of lower dentition of G. browni. 40 Plot 6a-d Bivariate scatter graphs showing comparison of the studied 54 specimens with the type specimens. Plot 7 Shows the size variation of cheek teeth L. pentapotamiae. 55 Plot 8 Width/length ratio of studied specimens L. pentapotamiae. 55 Plot 9 a-e Variation in the dentition of various species of Listriodon. 58 Plot 10 a-c Bivariate scatter graphs showing comparison of the studied 71 specimens with the type specimens. Plot 11 Width/length indexes of the cheek teeth D. majus. 72 Plot 12 Bivariate scatter graphs showing comparison of the studied specimens with the type specimens and previously studied 79 specimens. Plot 13a-b Bivariate scatter graphs showing comparison of the studied 89 specimens. Plot 14 Width/length indexes of cheek teeth of G. priscilla. 89 Plot 15a-j Bivariate scatter graphs showing comparison of the studied 124 specimens with the type specimens. Plot 16 Plot shows the size variation in the upper dentition of G. punjabiensis. 124

Plot 17 Plot shows the size variation in the lower dentition of G. 125 punjabiensis. xiii

Page Plot No. Title No. Plot 18 Width/length indexes of the upper cheek teeth of G. 125 punjabiensis Plot 19 Width/length indexes of the lower cheek teeth of G. 126 punjabiensis Plot 20a-c Bivariates showing comparison of the studied specimens 137 with the type specimens. Plot 21 Width/length indexes of the cheek teeth of Gazella sp. 138 Plot 22 Plot shows the size variation in the dentition of Gazella sp. 138 Plot 23a-c Bivariate scatter graphs showing comparison of the studied 148 specimens with the type specimens. Plot 24 Width/length indexes of the cheek teeth of Eotragus sp. 149 Plot 25 Plot shows the size variation in the lower dentition of 149 Eotragus sp. Plot 26 a-c Variation in the dentition of various species of genus 150 Eotragus. Plot 27 Comparison of the studied specimens with different species 151 of genus Eotragus.

1 INTRODUCTION

Geography of the Siwaliks Medlicot (1864) introduced the name Siwaliks for the Sub-Himalayan rocks and originally this term was derived from the Siwalik Hills in Deharadun (India). The term “Siwalik hills” was applied in a geographical sense by Falconer and Cautley (1849). It was from these Siwalik hills that the Upper Tertiary and Pleistocene fossil vertebrates were first discovered and studied by Hugh Falconer, hence the name “Siwalik” as applied to the bed containing the fossils. The Siwalik hills are situated within the political boundaries of Pakistan, India, Nepal and Bhutan, and their range is 6 to 90 km in width (Acharyya, 1994). They gradually become steeper and narrower in relief and width respectively, from northern Pakistan to Bhutan (over 2000 km in length, Figure1) (Chauhan and Gill, 2002; Khan et al., 2006). The Siwalik deposits are one of the most comprehensively studied fluvial sequences in the world. They comprise mudstones, sandstones, and coarsely bedded conglomerates laid down when the region was a vast basin during Middle Miocene to Upper Pleistocene times. The sediments were deposited by rivers flowing southwards from the Greater Himalayas, resulting in extensive multi- ordered drainage systems. Following this deposition, the sediments were uplifted through intense tectonic regimes (commencing in Upper Miocene times), subsequently resulting in a unique topographical entity, the Siwalik hills (Acharyya, 1994; Chauhan and Gill, 2002). The Siwalik hills form a ridge between the great flood plain of the Ganges River and the Himalaya Mountains. These hills are relatively low, ranging from three thousand to four thousand feet in height above sea level and they have a general northwest to southeast trend parallel to the Himalayas. They are distinguished by their asymmetric form, having in general a steep face or scarp on the southern side towards the valley and gentler slopes on the northern sides. The hills actually have their expression as a series of parallel ridges, forming a belt some eight miles or so in width. Behind and between these ridges are narrow valleys or dunes, running parallel to the general trend of the range and the largest of these, the Dehradun, separates the Siwalik Hills from the greater reaches of the Himalaya range. Many rivers traverse the Siwalik Hills, at right angles to their northwest and southeast trend, cutting through the highlands by narrow gorges and debouching on the plain to the south in sandy flats (Colbert, 1935).

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Figure 1: The Siwalik sediments along the foothills of Himalayas (Chauhan and Gill, 2002).

In Pakistan, Siwalik sediments are especially extending on the Potwar Plateau, where they are exposed in a folded belt extending from the Salt Range in the South to the Margalla hills in the North, and from the East of River Jehlem to the West of river the Indus (Pilbeam et al., 1979). The Potwar Plateau in the Punjab Province 72° 30' E 33o 00' N is an elevated area of 20,000 km2 and is constituted by a less internally deformed fold and thrust belt having a width of approximately 150 km in North-South direction. It is largely covered by the Siwalik sequence, though at places Upper Eocene Shale and limestone crop out locally in folded inliers. Its Northern part known as the North Potwar Deformed Zone (NPDZ) is more intensely deformed. It is characterized by East-West; tight and complex folds overturned to the South and sheared by steep angle faults, pop up structures and triangle zone. In the Western part this basin is comprised several East- Wests, broad and gentle folds. In the Eastern part, the strike abruptly changes to the northeast and the structures comprise tightly folded anticlines and broad synclines (Khan et al., 1986). GEOLOGY OF THE SIWALIKS The Siwalik Group comprises a thick sequence of terrestrial deposits in the northern part of the Subcontinent, with a relatively high degree of completeness over most of the Neogene’s time (Flynn, 2003; Flynn et al., 1995). The area of the Siwaliks, an elongated foreland basin of the Himalayas filled with molasse-type sediments of the Neogene and early Quaternary age, is developed at the foothills of the Himalayan 3 mountain belt. The stratigraphic sequence preserves a continuous record of the continental sedimentation (sediment thickness > 6 km at places) as well as an equally comparable continuous record of vertebrates, especially that of the mammals (Basu, 2004).

The Siwalik Group in Pakistan can be clearly divided, according to the lithological characters, into the usual three subgroups - Lower, Middle and Upper, and further into their formation scale of lithostratigraphic units. The Lower Siwaliks (Kamlial and Chinji formations) consists of a sequence of sandstone-mudstone couplets with a marked dominance of the mudstones over the sandstones. The development of paleosol horizons is also fairly frequent. The Middle Siwaliks (Nagri and Dhok Pathan formations) are dominantly arenaceous, consisting of multistoried coarse to medium-grained, blue- gray, massive sandstones (30 to > 60 m) with subordinate representation of clays, mudstones and siltstones. The Middle Siwalik sediments are known to produce scattered fossils. The Upper Siwalik subgroup is classified into three lithostratigraphic formations (Tatrot, Pinjor, Boulder Conglomerate formations). The three units comprise the sequences of the sandstone-mudstone couplets, the Parmandal Sandstone and the Boulder Conglomerate Formation, the upper most lithostratigraphic unit (Quade and Cerling, 1995). The Dhok Pathan Formation is composed of gray sandstone and red-brown mudstone with a few thin conglomerate interbeds. Sandstone and superposed red mudstone often form fining-upward couplets where the lower contact is erosional and lined with ripped-up clasts of the underlying mottled and red-brown clay stone. At a few places, thin crevasse-splay sheets, around 30 cm thick, clast-supported conglomerates occur. These conglomerate beds often contain unidentifiable bone and tooth fragments. Sandstone beds upper section gradually gets thicker as well as multistoried. These substantially thicker, vertically stacked and laterally extensive individual gray sandstone units form a fining-upward sequence with thinner dull red to brown siltstones on top. Varicolored, mottled, highly bioturbated paleosol horizons form distinct and laterally extensive units within the siltstone or at the transition of the sandstone to siltstone facies. In the lower Formation, there are pinkish to light brown hard, mottled, pseudo- nodular calcrete beds that are developed usually at the top of sandstone units, which are heavily bioturbated near the calcrete beds. Rhizoliths and worm burrows/trails are common in these hard carbonate-rich beds and lenses. The middle part of the Formation has alternating buff and light gray sandstone units, very much reminiscent of the Litra Formation of the Sulaiman Range (Behrensmeyer and Tauxe, 1982). Conglomerate 4 interbeds containing extra-formational pebbles of sandstone, quartzite, limestone and igneous rocks usually define the erosional base of the fining-upward sequences, but towards the upper section they form fairly thick laterally extensive independent bodies. The upper contact of the middle portion of the Formation is gradational, usually placed when the conglomerates frequently occur as massive to crudely bedded units, 1-3 m thick. These sand bodies also commonly have conglomerate lenses and scattered pebbles along the cross bed fore sets. The upper part of the Formation consists of thick massive to stratified conglomerate and pebbly gray sandstone with subordinate medium-to fine- grained friable gray sandstone and grayish-brown siltstone. Finer facies appear to be more common in the middle portion in the Bhandar area where they also contain some vertebrate fossils.

Lithology of the Siwaliks Siwalik lithology includes sandstones, siltstones, mudstones and rare marls and clays. The composition of the Siwalik beds show that they are nothing else than the alluvial detritus derived from the sub aerial waste of the mountains, swept down by their numerous rivers and streams and deposited at their foot. Siwalik rocks are the result of rapid and perhaps almost continuous stream erosion. Lithologically the origin of the series is probably simple, representing increasingly coarse river detritus brought down from a rapidly rising mountain mass. Moreover, there are no great secondary changes to be found in the Siwalik rocks, evidences of automorphism and glaciations being absent. The factor of predominant importance is that of erosion and deposition by rapidly flowing rivers. Paleosol sequences indicate reorganization of topography and drainage accompanying a transition to a more seasonal climate (Fatmi, 1973). Thus, the Potwar Plateau encompassed only a small part of this ancient foreland basin, providing only limited information on the whole system (Willis and Behrensmeyer, 1995). As a result of the deposition on alluvial fans and interfan areas, the sloping floodplains (Figure 2) formed a mosaic of low and high areas with local relief on the order of 10 m and at most a few hundred meters elevation difference across the floodplain (Willis, 1993a). Low areas were sites of short lived, perennial or seasonal ponds and swamps. Such low areas would have been rapidly filled by crevasse-splays and levees associated with the larger second and third order channels, and with the sediment from even smaller fourth-order floodplain streams (Figure 2). Preserved floodplain deposits therefore predominantly consist of low- 5 angle crevasse-splay lobes that build out into low areas, some of which contained wetlands or lakes, along with minor contributions from levees and small floodplain channels (Behrensmeyer et al., 1995; Willis and Behrensmeyer, 1995). A few paleosols may have been formed under water logged, grassy woodlands, but most were formed under drier conditions and more closed vegetation. The Miocene Siwaliks were deposited in a very large-scale fluvial system, one comparable in size to the modern Indus or Ganges systems. These modern rivers occupy divergent basins paralleling the northern and western mountains and bounded by the Indian craton to the south and east. This arrangement of mountains, basins and major drainages is a result of the tectonic movement of the subcontinent against Asia and existed throughout the Miocene. However, paleochannel directions and paleocurrents indicate that the paleo-Ganges system may have extended farther west draining the Potwar region (Beck and Burbank, 1990), which now is connected to the Indus drainage basin. The reconstructed Miocene Indo-Gangetic system extended over 2000 km to the east and 1000 km to the South, with floodplain widths on the order of 100 to 500 km. Reconstructions of the Siwalik rivers depict drainage basins coalescing into trunk rivers running to the Arabian Sea and Bay of Bengal along the axes of the two divergent basins. Large submarine sediment fans attest the existence of these Indus-size trunk rivers in the Miocene (Rea, 1992), but deposits of the axial system that drained the late Miocene Potwar are not preserved, either because it flowed eastward to the Bay of Bengal or, by flowing South to the Arabian Sea, it was confined by the slope of the floodplain to the margin of the depositional basin (Willis, 1993a, b; Willis and Behrensmeyer, 1995). However, the Channel and floodplain deposits of large and small tributaries to the Trunk Rivers are preserved. Comparison of the Chinji and Dhok Pathan formations indicates similarities in their depositional environments and in their transitional stratigraphic relationships with the intervening Nagri Formation (Behrensmeyer and Tauxe, 1982; Behrensmeyer, 1987; Willis, 1993b; Zaleha, 1994). The sediments of the Middle Siwaliks were deposited by coexisting fluvial systems, with the larger emergent of Nagri system followed by Dhok Pathan system. In comparison, Nagri floodplains were not well drained, with smaller rivers having more seasonally variable flow and more frequent avulsions. Both the Chinji and the Dhok Pathan formations have more well differentiated paleosols than the Nagri Formation, having overall differences in proportions of channel and overbank deposits. None of the formations have paleosols that are readily assigned to modern soil types, 6 although they are broadly similar to the modern soils of the Indo-Gangetic plain (Retallack, 1991; Behrensmeyer et al., 1995; Quade and Cerling, 1995), especially those forming under 25 C mean annual temperature and 1400 mm/yr precipitation (Behrensmeyer et al., 1995). Temporal changes in the Siwalik fluvial system have been documented, but the degree to which they are related to climate or subsidence, or to which they are simply due to the autocyclic dynamics of the fluvial system, is not clear. The transition between the Chinji and Nagri formations has been interpreted as due to the progradation of a second- order emergent system over a smaller, third-order interfan system (Willis, 1993a; Zaleha, 1997). The Nagri and Dhok Pathan transition, on the other hand, seems a well- documented case of the coexistence of two contemporaneous systems (Behrensmeyer and Tauxe, 1982). The Chinji floodplains were apparently more poorly drained than those of the Nagri Formation, with the abundance of lacustrine deposits and iron-concretion-rich paleosols indicating greater seasonal waterlogging. Floodplain deposition in the Chinji, and perhaps the Dhok Pathan Formation was also more episodic than in the Nagri, with longer-lived small floodplain channels. Initially, flow in these small channels was continuous year-round, but in later stages it became more episodic with periods of sub aerial exposure as the channels were abandoned and later infilled. By contrast, in the Nagri Formation even the initial stages of channel flow may have been more seasonal, with deposition of sediments occurring mostly during floods. Specific and important differences between the rivers of the Nagri and the Dhok Pathan formations include decreased channel size and discharge in the latter, as well as decreased avulsion period (Zaleha, 1997). 7

Figure 2: Reconstruction of Late Miocene Siwalik fluvial system in plan view. Numbers refer to orders of streams as discussed in text. 1= axial stem river, shown flowing eastward to the Bay of Bengal. 2= emergent or upland-sourced rivers, draining mountains and tributary to the axial stem river. 3= interfan or lowland-sourced streams, arising on the floodplain or near the mountain front and generally tributary to the axial or second-order rivers. 4= small streams of the floodplain, often ephemeral. Note that the third- and fourth-order streams may also have drained into low areas on the floodplain, creating seasonal swamps or ponds as indicated by gray stippled areas. The boxed inset approximates the size of the modern Potwar Plateau (Source, Barry et al., 2002). Sedimentology of the Siwaliks The Siwalik Miocene sediments are entirely fluvial in origin, having been deposited by large river systems. Some of the sections exceed 3000 m of such accumulated sediments and which are now exposed on the surface. These sedimentary rocks are usually divided into time successive formations, with the classic sequence of the Potwar comprising the Murree, Kamlial, Chinji, Nagri, and Dhok Pathan Formations of Pilgrim (1910, 1913) and what the Geological Survey of Pakistan refers to as the Soan Formation (Cheema et al., 1977). The important sedimentologic features of the rocks have been extensively discussed by Behrensmeyer (1987), Behrensmeyer et al. (1995), Willis and Behrensmeyer (1995), Stix (1982) and Willis (1993a, b). Because the sediments are well suited for magnetostratigraphic studies, it has been possible to establish a regional 8 chronostratigraphic framework that now spans the Early Miocene through the Pleistocene (Opdyke et al., 1979; Johnson, G. D., et al., 1982, Tauxe and Opdyke, 1982; Johnson, N. M., et al., 1982, 1985; Kappelman, 1986; Friedman et al., 1992; Downing et al., 1993; Ager, 1973). Changes in the rates of sediment accumulation and an apparent widespread depositional hiatus have made the stratigraphic relations and ages of the Early Pliocene formations of the Potwar less certain. These include the Tatrot Formation and its equivalents (Barry et al., 1982; Hussain et al., 1992) and some older sediment near Rhotas and Jalalpur. The high transport energy of running water in the river channels hardly permits the abundant deposition of small, light clay particles, except in specific environments like downstream alluvial plains and floodplains. Several workers (Bhattacharya and Misra, 1963; Bhattacharya, 1970; Chaudhri and Gill, 1983; Bagati and Kumar, 1994; Raiverman and Suresh, 1997; Raiverman, 2002; Biswas, 1994) have discussed the clay mineralogy of late Neogene sediments of the Middle Siwaliks. The first two drainages were major rivers draining through Higher and Lesser Himalaya, whereas the piedmont drainage, tributaries of Major River, drained through Sub-Himalayan region (Kumar et al., 1999; Ghosh et al., 2003). Interfingering of channel deposits was recognized on the basis of sand body geometry, colour, framework composition and paleoflow pattern. However, it is difficult to segregate the floodplain deposits of these different drainages due to no apparent colour difference among the mudstones. The Siwaliks has thick mass of fluvial deposits, comprising sandstones, mudstones and conglomerates of the Middle (Dhok Pathan Formation) and the Upper Siwalik (Tatrot, Pinjor and Boulder Conglomerate Formations) sub-groups. This sequence is characterized by stratigraphic coarsening up, stratal thickening and distinct changes in the channel body geometry and variation in the percentage of overbank to channel deposits. The basal stratigraphic succession (400 m) (prior to 5.5 myr), dominated by multistory gray sheet sandstones with copious erosional surfaces and trivial overbank mudstone. It lacks lateral accrual surfaces and low paleocurrent variability. The sizes of cross-bedded set (up to 2 m) and bank-derived intraclast lags indicate braided river system. Southeast paleoflow suggests the palaeodrainage was parallel to present Himalayan trend. Hence it represents the axial stem drainage system (Kumar et al., 1999). Between 400 and 600 m (5.5 and 5 Ma), the abundance of mudstone increased while the size of the sandstone bodies decreased. 9 Sediments of Himalayan foreland basin and contemporary Bengal Fan have high proportion of smectite during late Neogene (7.4 –0.5 Ma). Despite the escalation of monsoon in the Himalayan foreland basin at about 8 Ma, Quade et al. (1989) and Burbank et al. (1993). The variation in climatic conditions in the source area may have resulted in accumulation of different suites of clay minerals. Illite and chlorite formation predominate during less hydrolysing, cold-dry glacial periods, whereas smectite and kaolinite predominate during more hydrolysing warm-humid conditions during interglacial stages.

TAPHONOMY OF THE SIWALIKS Taphonomic research to date in the Siwaliks has focused on the distribution of fossil localities among depositional environments, inferring to conditions of mortality and accumulation, and reconstructing the profusion of taxa in the original community (Badgley and Behrensmeyer, 1980; Badgley, 1986; Raza, 1983; Behrensmeyer, 1988). The basis for assessing the quality of the fossil record and the nature of evolutionary and ecological questions that can be addressed for the analyses with the available data is provided by the taphonomic features of the fossil assemblages. Differences in the lateral distribution of sedimentary facies, in associated habitats and processes of mortality, and in post-mortem alteration, transport, and burial may considerably change the quantity, quality and identifiability of the preserved material. At the level of depositional system, the particular distribution of local environments determines the habitats available to organisms. The rates and processes of preservation may vary greatly among these environments in relation to biotic and abiotic components. Recognition of changes in taphonomic selectivity facilitates the distinction between apparent and real changes in original biotas (Koch, 1987; Badamgarav and Gingerich. 1988). Three vital aspects of fossil assemblages verify the reliability of inferences regarding the original faunal composition, the associations amongst taxa, the rates of morphological evolution within lineages and the patterns of immigration and extinction. Changes in the preservational bias may expose significant environmental changes that can be correlated with changes in biotic composition or fossil productivity (e.g. Badgley et. al., 1995; Behrensmeyer, 1988; Kidwell, 1988). Fossil assemblages from the Siwalik deposits reveal features indicative of fluvial transportation and deposition of abraded bones, bones dispersed through the sediment matrix, absence of skeletal association of the fossilized animals and lack of the more 10 transportable elements such as vertebrae and ribs. Teeth and jaws bones and fragmented bones are the major constituents of the assemblages. The taphonomic study of the fossil material reveal a variety of pre-burial and post-burial processes affected the bones and teeth deposited in the Siwaliks. Significant modifications were observed in the vast majority of the examined specimens. Extensive weathering cracks are indicative of the long-term exposure of the collected specimens on ground. Partly articulated, partly associated and mostly dispersed skeletal parts point out the long transportation and the significant dispersal of the occurred skeletal elements. Seismoturbation and faulting caused the post burial fracturing of various skeletal elements.

11 Siwalik Stratigraphy The Mio-Pliocene Siwalik group was deposited in the Himalayan foreland basin in response to uplift and erosion in the Himalayan fold-thrust belt. This 7 km thick succession of fluvial deposit is rich in vertebrate fauna. The Siwalik fauna has been regarded as one of the most important Cenozoic fossils anywhere in the world and has been intensively considered for more than a century. Several International and Pakistani scientists did post-independence geological work in the Siwaliks of Pakistan. As a result, a fine-scale magnetostratigraphic work has emerged, supplemented by vertebrate faunal zone. The important sedimentologic taphonomic features of the rocks have been extensively discussed by Behrensmeyer (1987), Behrensmeyer et al. (1995), Willis and Behrensmeyer (1995). The Siwalik sedimentary rocks are usually divided into time successive formations, with the classic sequence of the Potwar comprising the Murree, Kamlial, Chinji, Nagri and Dhok Pathan formations of Pilgrim (1910, 1913) and what the Geological Survey of Pakistan refers to as the Soan Formation (Cheema et al., 1977). The Chinji Formation is mud dominated while the Dhok Pathan Formation is sand dominated. It is often difficult to delineate the boundaries between the formations, however, form the geological or the sedimentological perspective it is best to view the Siwalik sequence as a single genetic unit. Nevertheless, the Siwalik formations have always been cryptic chronostratigraphic units and from the paleontological point of view, recognition of the formations and their boundaries has been a crucial step in dating the fossils (Colbert, 1935). This practice has, in the past, led to much confusion, but more recently magnetostratigraphy has provided more accurate dating of the rock. Pilgrim (1910, 1913) first recognized a series of successive “faunal zones,” initially using the term in a manner comparable to modern usage of the “stage” concept. Pilgrim’s units (Kamlial, Chinji, Nagri and Dhok Pathan) were bases on a combination of contained fauna and lithological criteria. In most instances their super positional relationships could be demonstrated, but the boundaries of the faunal zones were not delineated and, because or unreliable correlation, the faunal content of some zones could never be adequately differentiated. Subsequently, as stratigraphic concepts and nomenclature became more precise, Pilgrim’s faunal zones came to be used primarily either as lithostratigraphic formations or as chronostratigraphic “stages” or a combination of the two (Pilbeam et al., 1977; Barry et al., 1980, 1985; Flynn, 1986). Occasionally, they were recognized as being essential biostratigaphic units. Barry et al. (1980, 1982) first advocated in restricting 12 Pilgrim’s terms to the lighostratigraphic formations and later (Barry and Flynn, 1990) proposed a new series of biostratigraphic zones, called “faunal zones” in order to replace the Middle and Upper Siwaliks. Paleosol sequences indicate reorganization of topography and drainage accompanying a transition to a more seasonal climate. A few paleosols may have been formed under water logged, grassy woodlands, but most were formed under drier conditions and more closed vegetation. The Miocene Siwaliks were deposited in a very large-scale fluvial system, one comparable in size to the modern Indus or Ganges systems. These modern rivers occupy divergent basins paralleling the northern and western mountains and bounded by the Indian craton to the south and the east. This arrangement of mountains, basins and major drainages is a result of the tectonic movement of the subcontinent against Asia and existed throughout the Miocene. However, paleochannel directions and paleocurrents indicate that the paleo-Ganges system may have extended father west draining the Potwar region (Beck and Burbank, 1990), which now is connected to the Indus drainage basin. The reconstructed Miocene Indo-Gangetic system extended over 2000 km to the east and 1000 km to the south, with floodplain widths on the order of 100 to 500 km. Thus the Potwar Paltea encompassed only a small part of this part of this ancient foreland basin, providing only limited information on the whole system (Willis and Behrensmeyer, 1995). Reconstructions of the Siwalik Rivers depict drainage basins coalescing into Trunk Rivers running to the Arabian Sea and Bay of Bengal along the axes of the two divergent basins. Large submarine sediment fans attest the existence of these Indus-size trunk rivers in the Miocene (Rea, 1992), but deposits of the axial system that drained the late Miocene Potwar basin are not preserved, either because it flowed eastward to the Bay of Bengal or, by flowing South to the Arabian Sea, it was confined by the slop of the floodplain to the margin of the depositional basin (Willis, 1993a b; Wills and Behrensmeyer, 1995). However, the Channel and floodplain deposits of large and small tributaries to the Trunk Rivers are preserved. The Nagri and Dhok Pathan transition, on the other hand, seems a well-documented case of coexistence of the two contemporaneous systems (Behrensmeyer and Tausxe, 1982). The Chinji floodplains were apparently more poorly drained than those of the Nagri Formation, with the abundance of lacustrine deposits and iron-concretion-rich paleosols indicating greater seasonal waterlogging. Floodplains deposition in the Chinji, and perhaps the Dhok Pathan Formations were also more episodic than in the Nagri, with longer-lived small floodplains 13 channels. Initially, flow in these small channels was continues year-round, but in later stages it became more episodic with periods of sub-aerial exposure as the channels were abandoned and later infilled. By contrast, in the Nagri Formation even the initial stages of channels flow may have been more seasonal, with deposition of sediments occurring mostly during floods. Specific and important differences between the rivers of the Nagri and Dhok Pathan Formation include decreases channel size and discharge in the latter, as well as decreased avulsion period (Zaleha, 1997). Stratigraphic sections of the Siwaliks of Pakistan is described below (Figure 3).

Figure 3: Map of the Potwar Plateau (northern Pakistan) and a generalized stratigraphic section of the major Siwalik formations showing succession and ages (source Behrensmeyer and Barry, 2005).

14 Siwalik Chronostratigraphy The fossiliferous Neogene rocks of the Siwaliks are singular in their degree of completeness and represent almost the entire Neogene from about 22 Ma to less than 2 Ma (Keller et al., 1977; Opdyke et al., 1979; Azzaroli and Napoleone, 1982; Tandon et al., 1984; Johnson et al., 1985; Friedman et al., 1992). An extensive set of paleomagnetic determinations has provided good correlation within and between the various Siwalik sequences, between the Siwaliks through the GRTS, global record. The Potwar Plateau, and indeed the entire regional Neogene sequence as recorded in the Indo-Pakistan subcontinent, is apparently relatively homogenous and shows little, if any large patchiness. Thus, broadly contemporaneous faunas are basically similar throughout the region. Thus, it is difficult to know how closely a known stratigraphic range approximates the true stratigraphic ranges for all but the most common taxa. In this regard, Pilbeam et al. (1996) have conducted intensive biostratigraphic surveys in order to carefully document the first or last occurrences of a few common taxa. On the other hand, there are correlation problems created by several factors. Given the ages for the boundaries in the time scale used (Barndt, 1977; Berggren et al., 1985; Cande and Kent, 1995) and the accumulation rates; the calculated date for the “Hipparion” Datum in the Siwaliks is 10.7 Ma. This new date replaces previous estimates of approximately 9.5 Ma (Barry et al., 1982; Barry and Flynn, 1989), and like those dates, is based on stratigraphically older occurrences of this taxon than those used by Badgley (1986), who interpolated an age of 9.2 Ma. It is of more than historical interest that this “date” has varied with changes in the GRTS. For example, using the same stratigraphic occurrences under the Mankinen and Dalrymple (1979) time scale the date for the “Hipparion” Datum in the Siwaliks was 9.5 Ma, while with Berggren et al. (1985) it was 10.1 Ma and later with Cande and Kent (1995) it became 10.5 Ma. Barry et al. (2002) have published a comprehensive paper on Late Miocene of Northern Pakistan in which they developed a chronostratigraphic framework of the Siwaliks. Their study is based on 20 measured sections that range between 250 and 3200 m thick, as well as 27 shorter sections. 19 of the 20 long sections have determined magnetic-polarity stratigraphies and 16 of the shorter have at least some magnetic determinations. Apart from the two sections at Rohtas and Jalalpur (Opdyke et al., 1979; Johnson et al., 1982), the included sections form three regional networks: one on the northern limb of the Soan Synclinorium at Khaur, another on the Southern limb near Chinji, and a third at the Eastern end of the Potwar Plateau near Hasnot. However, 15 because of the distance and absence of continuous linkage exposures between regions, chronostratigraphic correlations between the three areas depend on the magnetic-polarity stratigraphy. No sections were correlated on the basis of biostratigraphy. In the Siwaliks individual stratigraphic sections of sufficient thickness that contain six or seven magnetic transitions (on the Potwar typically about 250 m) can usually be correlated to the Geomagnetic Polarity Time Scale (GPTS) (Johnson and McGee, 1983; Tauxe and Badgley, 1988). Although sections recording fewer transitions normally cannot be independently correlated to the GPTS, they can still be confidently placed in the chronostratigraphic framework by determining their stratigraphic relationships to the longer and better-constrained sections. This has been done by tracing isochronous lithological horizons laterally, such as sandstone bodies or paleosol horizons (Behrensmeyer and Tauxe 1982; Johnson et al., 1988; Badgley and Tauxe, 1990; Kappelman et al., 1991). The history of the Siwalik fauna is made up of seven episodes of faunal changes which are dated as older than 18 million years, 14-13 million years, 12 million years, 9.5 million years, 7.4 million years, with less chronologic accuracy, somewhat similar lithologic and faunal changes have been recognized in the Manchar, with less chronologic accuracy, somewhat similar lithologic and faunal changes have been recognized in the Manchar Formation of the Kirthar Province (coeval of Siwalik Group of Kohat Potwar Province) in south Indus basin (Barry and Flynn, 1989).

16 Siwalik Biostratigrapy and Paleoenvironments A major break-through providing an independent means of establishing ages of various local stratigraphic sections and correlation between different areas in the Siwaliks is obtained by applying magnetic-polarity, stratigraphy and fission track radiometric dating. A series of biostratigraphic zones, defined and characterized in fossiliferous, well- documented reference sections and correlated to magnetic polarity sequence, are believed to provide a more reliable framework for biostratigraphic division of the Siwalik faunas and should replace Pilgrim's "faunal zones" (Barry et al., 1982). A number of local stratigraphic sections in the Siwaliks of Potwar Plateau and their faunal and temporal correlation provides basis for a composite stratigraphic section, which then is designated as the Reference Section for the particular area. The biostratigraphic units summarized by Shah et al. (1985) as shown in Table 1 are, the biostratigraphic interval zone where each unit is defined as the stratigraphic interval between two distinctive biostratigraphic units recognized in the Reference Sections, and the base of each succeeding one is the top of the proceeding ones. Seven biostratigraphic interval zones along with of their characteristics fauna are given in Table 1.

Table 1. Biostratigraphic interval-zones scheme with characteristics fauna of the Siwalik Group (and Murree Formation) of the Potwar Plateau (Shah et al., 1985).

Time Biostratigraphic Chacteristic Fauna (Ma BP) Interval-Zone

1_ ------(1.5)------2 Elephus planifrons Elephas planifrons, Equus sivalensis, Hypolryus sp., Sivutherium giganteum, Cervids. 3_ ------(3.9)------4_ Hexaprotodon Hexaprotodon sivalensis, Proamphibos sivalensis lachrymans, Stegodon sp., Potamochoerus sp., Dorcatherium sp., Hippopotamodon sivalense, Percrocuta grandis. 5_ 17 ------(5.3)------6_ Selenoportax Mus auctor, Protuchyoryctes talroli, lydekkeri Selenoportax lydekkeri, Presbytis sivalensis, Induretos punjabiensis, Hipparion s.l., Hominoidea. 7_ ------(7.4)------8_ Hipparion s.l. Hipparion s.l., Progonomys spp., Hippopotamodon sivalensis, Propotamochoerus hysudricus, Sivahyus punjabienses, Dorcabune nagrii, Dorcatherium majus, Tetracondon magnus, Hominoidea. 9_ ------(9.5)------10_ 11_ 12_ Giraffokeryx Giraffokeryx punjabiensis, Antemus punjabiensis chinjiensis, Kanisamys potwarensis, Helicoporulas praecos, Sivoreas eremite, Miotragocerus glutens, Percrocuta carnifex, Doracatherium minus, Dorcqabune anthracothrioids, Hominoidea (Sivapithecus indicus), Listriodon pentapotamiae, Conohyus sindiensis. 13_

14_ ------(14.0)------15_ Listriodon sp. Listriodon spp., Conohyus spp., Eotragus, Prokanisamys sp., “Listridon sp.,” Kanisamys indicus, Sayimys sp., Sanitherium sp., Nimravidae. Small hominoid, Dorcabune sp., Dorcatherium sp. 18 16_ ------(16.3)------17_ 18_ Gomphotherium s.l. “Gomphotherium s.l.” Deinotherium pentapotamiae, Brachypotherium sp., gigantic anthracothere. 19_ 20_ 21_

Paleoenvironments of the Siwalik indicate two major environmental turnover as the older one is Asian monsoonal precipitation system and the younger one is transition from C3 floras to C4 grasses. The timing of monsoonal inception is controversial. According to Molnar et al. (1993) it estimates at 7.0 to 8.0 Ma while according to Kroon et al. (1991) it estimates at 9.0 to 11.0 Ma. The transition from C3 dominated to C4 dominated vegetation is documented approximately 8.1 Ma and with fully C4 floras at 7.4

Ma with last occurrence of C3 floras at 7.0 Ma. This transition had a marked effect on mammalian feeding ecology. Different taxa described from Siwaliks indicate that the separation of European and South Asian biogeographic thearers occurred at middle Miocene (Barry et al., 2002). The thick fluvial and lacustrine environments of the different formation of the Siwalik Hills of Pakistan indicate the Presence of lakes and rivers systems during the depositional stages (Willis 1993a, b; and Behrensmeyer and Tauxe, 1982). Permanent forests and woodlands with some interspersed grasses (mostly

C3) were present about 9 Ma, after which wooded grasslands became widespread on floodplains (Quade et al., 1989; Morgan et al., 1994).

19

STUDIED SECTION Dhok Bun Ameer Khatoon village is located in the district Chakwal, Punjab, northern Pakistan. The village (32o 47' 26.4" N, 72° 55' 35.7" E) is surrounded by Miocene deposits of the Lower Siwaliks (Figure 4). It is not only rich in fossil vertebrates but also in unique colored shale, containing some amount of unweathered igneous minerals, notable feldspar. It is composed of red brown mudstone with common grey sandstone inter-beds. The Dhok Bun Ameer Khatoon fauna mainly consists of Artiodactyla (suids, tragulids, giraffids, cervids and bovids) and Perissodactyla (rhinoceros). Giraffids are more abundant than those of the other taxa (Plot 1 and Plot 2, Appendix).

Figure 4: Map of the studied section Dhok Bun Ameer Khatoon, district Chakwal, Punjab, Pakistan. 20

Dhok Bun Ameer Khatoon fauna

3

2

No. of species 1

0 Rhinocerotidae Suidae Tragulidae Giraffidae Bovidae Family

Plot 1: Dhok Bun Ameer Khatoon even and odd-toed families and number of species described in this thesis.

Plot 2: Dhok Bun Ameer Khatoon even and odd-toed species and number of specimens described in this thesis. Methodology The Miocene hills of the Dhok Bun Ameer Khatoon were investigated thoroughly. Numbers of field trips were carried out to the studied section by 2006 and 2007 and 21 different collecting methods were employed for the collection of fossils. Surface collection has been the primary mean of collecting fossil remains. Excavations were also conducted at some places of the locality where dense concentrations of fossil bones occur in situ within sandstone. The embedded material was carefully excavated with the help of chisels; geological hammers fine needles, penknives, hand lenses and brushes. Then this collected material was transported to the laboratory. As a result valuable and worth identifying specimens of mammalian fossils were discovered. Among them even and odd- toed ungulates were dominant. In the laboratory, the material was carefully washed, cleaned, and prepared for the taxonomic study. The broken parts were assembled by using various types of gums (resins) such as Araldite, Peligom, Elfy, Elite, Magic stone and Fixin. The catalogue number of the PUPC specimens consists of series, i.e. yearly catalogued number and serial catalogued number, so figures of the specimen represent the collection year (numerator) and serial number (denominator) of that year (e.g. 08/100). All measurements are given in mm, with an accuracy of one decimal digit. The dental length (l) was measured on the occlusal surface. The tooth width (w) is the maximum width. Comparisons were made with specimens from the Natural History Museum, London (BMNH), the American Museum of Natural History (AMNH), the Geological Survey of Pakistan (GSP), the Geological Survey of India (GSI), and the specimens from the Palaeontology laboratory of the Zoology department of the Punjab University, Lahore, Pakistan (PUPC). The studied material is stored in the Palaeontology laboratory of the Zoology Department of the Punjab University, in Lahore, Pakistan.

22 TOOTH MORPHOLOGY Tooth cusp nomenclature in this thesis follows that of Akhtar (1992), Janis and Scott (1987a, b), Gentry (1994), Heissig (1972) and Cerdeno (1995) as shown in the figures 5 A-D. An entostyle can be founded in the center of the lingual side of the upper molar and ectostylid is found in the buccal side of the lower molar, completely or partly separate from the rest of the occlusal surface.

23

Figure 5 A-D: Terminology of tooth crown elements in even and odd toed mammals. A: Upper third premolar. B: Lower third molar. C: Lower fourth premolar. (Akhtar, 1992; Janis and Scott, 1987 a, b and Gentry, 1994). D: Rhinoceros lower molar modified after Heissig (1972) and Cerdeno (1995).

24 Objectives of Present Study The main aim of this study is to provide the first complete citation of the even and odd-toed mammalian fossils found in the vicinity of the Dhok Bun Ameer Khatoon by highlighting the aspects of taxonomy and paleontology of the Siwaliks of Pakistan. An important group, the even and odd-toed mammals were selected for the study as the collected material presented have notable diversity and thus can provide significant ecological, taxonomical, biostratigraphic and palaeoenvironmental information.

THESIS LAYOUT

This thesis consists of four distinct, separate and autonomous thematic units structured in a format that will be directed by Doctoral Programme Coordination Committee of the University of the Punjab, Lahore, Pakistan. Consequently, repetition of description, discussion and systematic about the genera does occur. The first chapter entitled “Introduction” includes mainly geography, geology, lithology, sedimentology, stratigraphy, biostratigraphy, and chronostratigraphy taphonomy and dental terminology. The second chapter is entitled “Review of Literature”. In the third chapter entitled “Systematic Palaeontology” taxonomical and morphological features of the studied material is elaborated and presented. The fourth chapter includes discussion and conclusion. The references and appendix are given at the end of the thesis. 25 LITERATURE REVIEW

An extensive work has been done on the Siwaliks by national and international teams of the palaeontologist (Falconer and Cautley 1836, 1849; Lydekker, 1876, 1882, 1883a, b, 1884, 1886; Pilgrim, 1910, 1912, 1913, 1937, 1939; Bakr, 1969, 1986; Sarwar, 1977; Akhtar, 1995, 1996; Khan et al., 2006, 2007, 2008 & 2009 and Metais et al., 2009). Recently a new species of the genus Microbunodon (Artiodactyla) has been described from the Miocene of Pakistan (Lihoreau et al., 2004). Welcomme et al. (2001) reported terrestrial detrital facies from the Bugti Hills in the South-Western Sulaiman geological province (Balochistan, Pakistan). This region has yielded the richest Tertiary vertebrate faunas to be found in Asia thus far. Barry et al. (2002), Cheema (2003), Flynn (2003) and Lihoreau et al. (2004) published comprehensive papers on the Siwalik mammals and the palaeoenvironments of the Siwalik deposits of the Potwar Plateau. The large mammalian Siwalik fauna has been the focus of many researchers (Savage and Russell, 1983; Nanda and Shani, 1990; Scott et al., 1999; Metais et al., 2000, 2001, 2004, 2009; Geraads et al., 2002; Barry et al., 2002, 2005; Bernor et al., 2003; Kaiser, 2003; Kaiser et al., 2003; Kaiser and Fortelius, 2003; Raymond et al., 2004; Behrensmeyer and Barry, 2005). Barry et al. (2002) in their comprehensive paper investigated faunal and environmental change in the Late Miocene Siwaliks of the northern Pakistan and described straitigraphic ranges of the fossil fauna (Appendix-4). Metais et al. (2003) reported dental and postcranial material of the bovid-like new artiodactyl ruminant mammal Palaeohypsodontus zinensis from the Late Oligocene of the Bugti Hills (Balochistan, Pakistan). This finding extends the geographic distribution of Palaeohypsodontus which was previously restricted to the early Oligocene of Mongolia and China. Barry et al. (2005) reported Late Oligocene and Early Miocene fossil ruminants from Nepal and Moroto in Uganda and the Zinda Pir sequence in Pakistan, which provide evidence for the existence of diverse latest Oligocene and earliest Miocene fossil faunas in Sub-Saharan Africa and Southern Asia. In addition, the Siwalik deposits of Pakistan yield numerous superposed assemblages that record the small mammal fauna throughout the Middle and Late Miocene of the Subcontinent (Flynn and Morgan, 2005). Brief background knowledge about the studied taxa is mentioned below: Rhinocerotids The Oligocene and Miocene deposits of Dera Bughti, Baluchistan, Pakistan yield various rhinocerotid genera (Foster-cooper, 1934; Antoine and Welcome, 2000; Antoine 26 et al., 2004; Metais et al., 2009). Metais et al. (2006) reported a fossil material of the enigmatic cetartiodactyl Bugtitherium grandincisivum from the Late Oligocene of the Bugti Member of the Chitarwata Formation in the Bugti Hills, Balochistan, Pakistan. The Miocene sediments of Africa, China, North America, Europe and Pakistan yield large mammalian fossils especially rhinoceroses (Hooijer, 1963; Savage, 1967; Heissig, 1969, 1972, 1999; Sarwar, 1971; Hamilton, 1973; Aguirre and Guerin, 1974; Ginsberg, 1974; Solounias, 1981; Qi and Yan, 1982; Zhang, 1982; Grooves, 1983; Savage and Russel, 1983; Brunet et al., 1987; Geraads, 1988; Xu, 1989; Wang, 1992; Cerdeno, 1995, 1996, 1998; Prothero, 1989; Holbrook, 1999; Cerdeno and Sanchez, 2000. Deng (2004, 2006) reported some species of the rhinoceroses in China that appears in the more recent levels of the Chinji Formation. Deng (2007) also described a skull of Parelasmotherium, a giant elasmotherine rhinocerotid with a huge nasal horn, from an Early Late Miocene locality of the Linxia Basin in Gansu, China. Nevertheless the systematics of the Oligocene and Miocene rhinoceroses is confused. Many generic names have been used besides those do far mentioned while multitudes of species level names have been founded and used in differing combinations with the generic names. Colbert (1942) compared the cranial and dental characteristics of Rhinoceros sondaicus with Gaindatherium and suggested that Rhinoceros sondaicus is morphologically primitive among extinct and extant Rhinoceros although its remains have been recovered from the Middle and Late Pleistocene of Asia. At present, the transition from the Gaindatherium lineage to the Pleistocene Rhinoceros species is poorly known (Heissig, 1989). Moreover, their earliest fossil remains of Rhinoceros sondaicus had previously been recorded from the middle Pleistocene Djetis Bed and Trinil Bed of Java (Hooijer, 1957). Therefore, the discovery of the Rhinoceros sondaicus from the Early Pleistocene of Myanmar fills the geological and chronological gap between primitive Gaindatherium and the Middle Pleistocene Rhinoceros sondaicus from Java. This discovery suggests Rhinoceros sondaicus originated as early as the Early Pleistocene in continental Asia, and its possible migration to island Southeast Asia during the Late Early Pleistocene and later ages (Thein et al., 2006).

Suids The Suoidea (pig-like animals) is a superfamily of the Artiodactyla (even-toed mammals), which is further divided into four families Suidae (pigs), Tayassuidae (peccaries), Sanitheriidae (extinct) and Hippopotamidae (hippopotami) (McKenna and 27 Bell, 1997). The sister relationship between Suidae and Tayassuidae is attested to on both morphological (Gentry and Hooker, 1988) and molecular basis (Irwin and Arnason, 1994; Randi et al., 1996), but the closer relationships of Hippopotamidae and Sanitheriidae with suoids are in doubt. Van der Made (1999) raised the Hippopotamidae to the superfamily Hippopotamoidea and moved it out of Suoidea, supporting the opinion of a closer relationship between hippopotami and anthracotherids (Colbert, 1935; Viret, 1961; Gentry and Hooker, 1988). Pigs and peccaries inhabit similar adaptive zones (Simpson, 1944) in the Old and New World, respectively. The suoids are primarily omnivorous, and seem to be the most primitive living artiodactyls, retaining low-crowned cheek teeth with simple bunodont cusps, four distinct digits, separated foot bones, absence of frontal appendages, and a simple, non-ruminating stomach. Unlike the highly diverse ruminants, the suoids today form a small and simple group of artiodactyls, with only seven genera and 12 species worldwide (Liu, 2003). Listriodon suid is documented on Europe, Africa, and Asia (Dong, 1987). The earliest European and African records of Listriodon are from early Middle Miocene (Koenigswald, 1967). There are two separate branches of Listriodont pigs that probably divided in the course of the MN4 unit. The branch that has become lophodont is placed in Listriodon, while the branch that remained sublophodont is placed in Bunolistriodon (Stefanovic, 2004). Simpson (1945) has categorized the Listriodontines on the basis of age into a single genus that belongs to Neogene. Debating on the validity of the genus Bunolistriodon, Gabunia (1973), Leinders (1975), Pickford (1986) and Pickford and Morales (2003) are of the view that Bunolistriodon Arambourg, 1963 and Listriodon Von Meyer, 1846 should be merged into the single genus. The origin of listriodonts is considered to be from Eurasia from where they spread to Africa. During MN4 they underwent dental morphological changes from bunodont, becoming lophodont reaching Africa in MN6. During MN6 Asian suid Listriodon entered into Europe (Thenius, 1952, Made, 1989-1990). They are known immigrants to the region and may represent relatively abrupt immigration events (Fortelius and Jernvall, 2004). This took a whole time of almost 4-5 million years. They were not found in Europe, Africa, India and China at the end of Late Miocene. Some discoveries show their presence in the later MN successions but they became extinct after the Lower Pliocene in Europe, with not a single appearance in MN9 (Pickford and Morales, 2003). In Asia the last occurrence extends into Late Pliocene. 28 In Sub-continent, Listriodon has been described by several workers from Siwalik region. The specimens discovered from Chinji belong to MN6-MN7/8 as is confirmed from the biostratigraphy and palaeomagnetic stratigraphy of the region (Pickford and Morales, 2003). There are strong similarities between Miocene Suid faunas of Europe and the Sub-Himalaya, especially among Listriodontinae. On the basis of these similarities this concept is refused that India was the centre of origin and diversification of the family Suidae (Pickford, 1988).

Tragulids Tragulids have occurred in both Eurasia and North America since the Middle Eocene (Vislobokova, 2001). The earliest tragulids, dependent on stable warm forests, are known in East Africa, Pakistan, and Europe by 18.0 million years (Gentry, 2000). Three groups, archaeomerycids, lophiomerycids, and bachitheriids, seem to exist only in Eurasia. Gelocids and tragulids occurred in Eurasia and Africa respectively. In America, tragulids were represented by leptomerycids and gelocids. Hypertragulids, a mainly North American group, were spread from the late Middle Eocene through to the beginning of the Oligocene in both the Old and New Worlds and up to the Early Miocene in North America. Although, there is no direct evidence on the place of origin of tragulines, Asia seems to be very plausible as the site of their early development. The primitive structure and a large diversity of Eocene tragulines in Asia also confirm the Asian origin of this group (Vislobokova, 1997). Several traguline genera occurred in the Middle Eocene of Asia. The most ancient of them were represented by the archaeomerycids Archaeomeryx and Xinjiangmeryx. Tragulids are traditionally considered as the most primitive living ruminants (Janis, 1984) and they still survive as tropical relicts: the water chevrotains of Africa and the mouse deer or Asiatic chevrotain of Southeast Asia. They are characterized by their skeletal and dental features, which are primitive within ruminants and their general shape, digestive system and ethology, which are reminiscent to those of pigs (Dubost, 1965). The tragulids are well known in the Siwaliks of Pakistan and they have been reported from the Chinji, Nagri and Dhok Pathan formations (Farooq, 2006). In the Chinji Formation, both genera of the Tragulidae family, Dorcatherium and Dorcabune are reported. Two species of Dorcatherium D. majus and D. minus are known from this Formation, whereas one species of Dorcabune D. anthracotherioides is reported. The Nagri Formation also reveals the two Siwalik genera Dorcatherium and Dorcabune. In 29 the richly fossiliferous Dhok Pathan Formation both genera have been reported (Khan and Farooq, 2006a). The extinct genus Dorcatherium is known from Asia (Lydekker, 1876; Matthew, 1929; Corbet and Hill, 1980), Europe and Africa. Until now many species of the genus Dorcatherium have been reported from Africa, Europe and Asia. The three Pakistani species of the genus Dorcatherium D. majus, D. minus and D. minimus are reported from the Siwaliks. The Tragulids entered Asia and Europe in the Late Miocene but have survived to date only in Asia, as they went extinct in Europe in the Late Pliocene. In Africa they first appeared in the Pleistocene and have lived there up to date. Climate became drier late in the Miocene and at about the same time (around 7.0 Ma) tragulids declined in the Siwaliks (Gentry, 2000).

Giraffids The Siwalik giraffid fauna was thoroughly described by Falconer and Cautley (1836); Falconer (1845); Forsyth Major (1891); Pilgrim (1911); Bohlin (1926); Colbert (1933, 1935a); Lewis (1939); Pincher (1949); Churcher (1978); Heintz et al. (1981); Akhtar et al. (1991); De Bonis et al. (1997) and Franz-Odendaal and Solounias (2004). They provide description of the main aspects and very complete account on different fossil species. Geraads (1979, 1985, 1986, 1989) described systematic and phylogeny of Miocene and Pleistocene giraffids from Greece, Macedoine and Baltimore. Godina (1997) gives new findings on Sivatherinae from the territory of the USSR. Hamilton (1978) explores giraffid remains from the Miocene of Africa as well as revised the phylogeny of the Giraffoidae. The Siwalik Pliocene and Pleistocene giraffids were discussed by Harris (1976, 1987a, b), Khan (1987) and Khan and Sarwar (2002). Late Miocene giraffids from Africa were described by Singer and Bone (1960), Pickford (1975) and Thomas (1984). Solounias and Moelleken (1993) and Solounias et al. (2000) analyse paleodiet and dietary adaptations of some extinct ruminants including giraffids by determining premaxillary shape. Geraads (1999) described a sub-adult skull from the Late Miocene of Kavakdere and referred it to the Indian genus Bramatherium, which increases the similarity between the Indian sub-continent and the Greco-Iranian province while Mitchell and Skinner (2003) reviewed the literature about the origin, phylogeny, and evolution of modern giraffes Giraffa camelopardalis. Franz-Odendaal, (2004) analyse dietary evaluations of an extinct giraffid, Sivatherium hendeyi from Langebaanweg, South Africa. 30 In the Siwaliks, the giraffids made their first appearance in the basal part of the Chinji Formation. The large sized giraffids (Sivatherine) were not present in the Chinji Formation. As indicated by the fossil record, the Sivatherines were definitely present in the Nagri Formation. Since the Nagri Formation, the record is highly fragmentary, and nothing can be stated with certainty about the quality of the group. Conversely, the Dhok Pathan was certainly the age of the gigantic giraffids. Many species of the Sivatherines are known from the beds of this age (Khan and Farooq, 2006a).

Bovids The bovids appear to have had three major adaptive radiations at 14, 7.5, and 2 Ma (Gentry, 1966, 1970, 1978; Solounias, 1982; Thomas, 1984; Ye, 1989), but few fossils are known from sediments older than 14 Ma (Solounias et al., 1995). Kingdon (1982) described phylogenetic diagram charting the evolution of the major bovid clades. Citations of very early Bovidae and the great diversity in the primitive forms in Central Asia, Mongolia and China and the sudden appearance of the bovids belonging to different subfamilies in Europe, the Indian Subcontinent and Africa, suggest that their origin can be traced in north or central Asia (Metias et al., 2001). The Miocene fossil record is both rich and geographically extensive, and as a result the reconstruction of the evolutionary history of Bovidae promises to provide one of the most complete such histories for any large mammal group (Bibi, 2009; Leslie and Koustubh, 2009). The earliest known boselaphine, if Eotragus can be excluded, may come from Pakistan (Solounias et al., 1995) at 17.6 Ma (Eotragus noyei). In Europe boselaphines only appeared in the later Middle Miocene. Miocene genera of boselaphines nearly all had keels on their horn cores. Around the end of the Upper Miocene Tragoportax vanished. In its place Bovini appeared in Eurasia, especially in India, and in Africa (Khan et al., 2009). Several species of gazelles were present during Late Miocene and Early Pliocene in Europe, Asia, and Africa (Savage and Russell, 1983). The widest distribution of Gazella probably occurred in Pliocene when it reached England (Heptner et al., 1988). The genus Gazella was erected by Blainville (1816) and was recorded for the first time as fossil horn-cores of G. stehlini (Gentry, 1966). Gazella is among the oldest of contemporary bovid genera and first appeared in the fossil record for the first time in the Rusingan (Early Miocene) of Libya. Gazella first appears in Europe (G. stehlini in Astaracian) and in Asia (Savage and Russell, 1983), during the Middle Miocene. Most species of Gazella from Europe are found as fragmentary material 31 such as isolated teeth and horn-cores and these species are small. G. capricornis, the best known of the European Pontain species is represented by more material than G. deperdita (Gentry, 1966). The progressive feature of G. lydekkeri is a hypsodonty. In the Namurungule Formation, Gazella are more hypsodont (Nakaya et al., 1984, 1987). The Chinese Pontain gazelles are more progressive especially the species G. lydekkeri although the width at the orbits is greater. The horn-cores seem to be identical in their morphology with G. lydekkeri and the nasal are somewhat shorter. In G. dorcadoides and G. altidens the teeth are still more hypsodont than G. lydekkeri (Khan et al., 2009). Gazella is recorded from the Lower and the Middle Siwaliks (Pilgrim, 1937; Akhtar, 1992; Khan, 2007, 2008). Pilgrim and Hopwood (1928) simply summarized all the nomenclature of fossil gazelles propagating the existence of a large number of species. G. lydekkeri recovered from the Late Miocene and the Middle Miocene assemblages of the Siwaliks respectively (Pilgrim, 1937, 1939). The occurrence of Gazella in Africa has been reported by Bourguignat (1870), Pomel (1895), Bate (1940), Arambourg (1947, 1957 and 1959), Dietrich (1950), and Gentry (1970). The record by Bourguignat (1870) is based upon his new species G. atlantica while that recorded by Bate (1940) from Palestine is based upon two species, G. decors and G. arista. Arambourg (1957) discovered a new species G. tingitana from the Late Pleistocene of Africa. Pomel (1895) described various species of Gazella from the Pleistocene of Algeria (Akhtar, 1992). G. gazella was the only gazelle species documented in Israel until after the Pre- Pottery Neolithic periods (Tchernov et al., 1986). In the last glacial period its range expanded southward, reaching the southern Sinai Peninsula. Gentry (1970) and Solounias (1981) proposed that the Chinese G. gaudryi was either conspecific with or in the same group as the contemporaneous European G. capricornis from Samos and Pikermi. The first female gazelles with horns probably occurred in Late Miocene, and this may have been related to their expansion into more open habitats (Gentry, 1990). Teilhard and Trassaert (1938) documented taxonomic difficulties in the genus Gazella from North China from the Yushe Basin. It is the first record of the genus in the Yushe region as well as in China and possesses characters that are more primitive than in any other member of the genus. Chen (1997) recognized seven species i.e. G. gaudryi, G. gaozhuangensis., G. yushensis, G. nihensis, G. blacki, G. sinensis, G. subgutturosa from the Neogene of the Yushe Region, three of which are new. G. gazella itself is not known from earlier than the Upper Pleistocene. Presumed Early Holocene occurrences are from 32 Karain Cave, Turkey, central Lebanon, Petra, Jordan, Hesban, east of Jericho and West Bank areas, where it has been described as G. gazella occurring with G. dorcas (Uerpmann, 1986). G. gaudryi is distinct from other contemporaneous “Early Pliocene” species in the world with its short and gracile horn-cores. G. blacki is distinct from contemporary members of the genus in Europe and Africa by its size, degree of hypsodonty, and structure of its P4, whereas the Pleistocene G. sinensis differs from all other members of the genus in its excessive cranial and horn-core robusticity (Chen, 1997). 33

SYSTEMATIC PALAEONTOLOGY

Class: MAMMALIA Linnaeus, 1758 Subclass: THERIIFORMES (Rowe, 1988) Mckenna and Bell, 1997 Order: PERISSODACTYLA Owen, 1848 Family: RHINOCEROTIDAE Owen, 1845 Subfamily: RHINOCEROTINAE Owen, 1845 Tribe: RHINOCEROTINI Owen, 1845 Subtribe: RHINOCEROTINA Owen, 1845

Genus: Gaindatherium Colbert, 1934 Type Species: Gaindatherium browni Colbert, 1934

Included species 1. G. browni Colbert, 1934 2. G. vidali Heissig, 1972

Diagnosis: An upper tertiary rhinoceros of medium size, with a saddle-shaped skull having a single horn on the nasals and with brachyodont, simple molar teeth. The orbit is located in an approximately central position above the first molar; the occiput is vertical; the postglenoid and posttympanic are fused, forming a closed tube for the external auditory meatus. There are two upper incisors, of which the lateral one is quite small; the upper molars are without an antecrochet or crista, and the crochet is but slightly developed. So the cheek teeth are brachydont and relatively simple, without antecrochet or crista, but with a crochet present in the last molar (Colbert, 1935).

Distribution: Lower to Middle Siwaliks.

Stratigraphic Level: The Chinji Formation.

34 Gaindatherium browni Colbert, 1934

Type Specimen: AMNH 19409, an almost complete skull.

Locality: Vicinity of Chinji Rest House, south of the Chinji village, Salt Range, Attock District, Punjab (Colbert, 1935).

Stratigraphic range: Lower Siwaliks, Chinji zone up to lower portion of the Middle Siwaliks.

Diagnoses: The specific diagnosis is the same as the generic diagnosis.

Specimens Examined: PUPC 08/14, an isolated right second lower premolar and PUPC

08/85, right mandibular ramus with M2-3.

Locality: Dhok Bun Ameer Khatoon, Chakwal district, Punjab, Pakistan.

Stratigraphic range: Lower Siwaliks (Chinji Formation).

Description

PUPC 08/14 (Figure 6) The specimen under study is an isolated right lower second premolar. It is finely preserved and in an early stage of wear. No trace of cement is present. The trigonid is V- shaped with the narrow and short paralophid and have right-angled metalophid. The talonid is U-shaped with the hypolophid and the entoconid. The labial groove is deep. The premolar is triangular in outline. The cingulum is absent. The paralophid is slightly shorter than the metalophid.

PUPC 08/85 (Figure 7) Mandible

The specimen under study is a broken right mandibular ramus with M2-3. It is damaged anteriorly and having the preserved ascending ramus which is 46 mm, antero- posterior length of bone is 155 mm, height below M2 is 55 mm and the thickness of bone 35 below M2 is 30.5 mm. Length of molar series is 75 mm. M2 and M 3 are well preserved while roots of P4 are also present and are not enough to show the morphological characters. Many details have been vanished because of major detoriation. The ramus is well preserved and the teeth are in middle wear. Thick enamel is present in both M2 and M3. The trigonid is V-shaped with the narrow and short paralophid and have right-angled metalophid. The talonid is U-shaped with the hypolophid and the entoconid. No trace of cement is present. There are neither lingual nor labial cingula. Posteriorly the ectolophid groove is marked to the base of the crown and is deep. The paralophid is present and crushed in M2, whereas in M3 it is completely preserved.

Hypolophid is oblique but transverse in occlusal view. In M2 the enamel is thick 2 mm, thinly wrinkled vertically and broken from anterior side of the molar. The measurements of the described teeth are provided in table 2. 36

A

B

C

Figure 6: G. browni, PUPC 08/14, an isolated lower right second premolar, A - buccal view, B - occlusal view, C - lingal view. Scale bar 10 mm. 37

A

B

C

Figure 7: G. browni, PUPC 08/85, a right mandibular ramus with M2-3, A - buccal view, B - occlusal view, C - lingal view. Scale bar 10 mm. 38

Table 2: Comparative Measurements (in mm) of teeth referred to G. browni Colbert, 1934. (Data taken from Colbert, 1935 and Hessig, 1972)* The studied specimens.

Species Number Position Length Width W/L

G. browni PUPC 08/14* P2 28 17 0.61

G. browni PUPC 08/85* M2 36 25 0.69

G. browni AMNH 29839 M2 43 28 0.65

G. browni PUPC 08/85* M3 39 24 0.62

G. browni PUPC 02/11 P2 28 19 0.68

G. browni AMNH 29838 P2 28.5 21.5 0.75

R(G) browni - P2 26 17 0.65

R(G) browni - M3 42 29 0.69

G. browni AMNH 29838 M3 43 26 0.60

G. browni PUPC 02/155 M3 44 23.5 0.53

39

Lower P2

25

20

15 PUPC 08/14* PUPC 02/11

Length 10 AMNH 29838 R(G) browni

5

0 25.5 26 26.5 27 27.5 28 28.5 29 Width

a

Lower M2

28.5 28

27.5 27 PUPC 08/85* 26.5 AMNH 29839 Length 26 25.5

25 24.5 35 36 37 38 39 40 41 42 43 44 Width

b

Lower M3

35

30

25 PUPC 08/85* 20 R(G) browni AMNH 29838

Length 15 PUPC 02/155 10

5

0 38 39 40 41 42 43 44 45 Width

c

Plot 3a-c: Bivariate scatter graphs showing comparison of the studied specimens with the type specimens. 40

W/L index between lower P2, M2-M3

0.8

0.7

0.6

0.5 P2 0.4 M2 M3 0.3

0.2

0.1

0 1 2 3 4

Plot 4: Width/length indexes of the cheek teeth of G. browni.

Lower P2-M3

35

30

PUPC 08/14* PUPC 08/85* 25

AMNH 29839 PUPC 08/85* 20 PUPC 02/11 AMNH 29838

Length 15 R(G) browni R(G) browni

10 AMNH 29838 PUPC 02/155

5

0 0 10 20 30 40 50 Width

Plot 5: Size variation of lower dentition of G. browni. 41 Discussion Colbert (1934) defined the genus and species Gaindatherium browni as a rhinoceros with several homologies with the extant species Rhinoceros sondaicus and R. unicornis and Heissig (1972) included it as a subgenus of Rhinoceros. Gaindatherium has been considered as a smaller genus when compared with R. sondaicus (Colbert, 1942). Gaindatherium has a middle-late Miocene distribution with two successive species i.e. G. browni and G. vidali described in the Siwaliks (Heissig, 1972; Sehgal and Nanda, 2002). G. browni share certain resemblances in the dental morphology with G. vidali from the Nagri Formation (Hessing, 1972). The resemblance lies in the presence of anterior posterior cingula, absence of lingual cingulum in the molars and absence of crista. However both the species have marked differences. G. vidali differ from the G. browni in having the well developed crochet, smaller size; well developed parastyle and parastyle fold, and funnel shaped postfossette. Dimensions of G. browni are larger than G. vidali. G. browni is known from the Lower Siwaliks (Chinji Formation) to the Middle Siwaliks (Nagri Formation) and other pre-hipparion localities of the Siwalik hills of Pakistan (Tang and Zong, 1987). According to Colbert (1935) Gaindatherium is a form more or less directly ancestral to the modern Asiatic rhinoceros and that it represent an intermediate link between the stem Caenopus type of true rhinocerine and the modern one horned rhinoceroses. The check teeth are brachyodont and relatively simple, without anterrochet or crista, but with a crochet present in the last molar. The present studied specimens have same character as mentioned for Gaindatherium by Colbert (1935). The present specimens when compared to Chilotherium intermedium were found smaller in size and different in morphology. The paralophid is very short and weak in P2 of C. intermedium, whereas paralophid in the specimen under study is short but thick as compared to C. intermedium. In the present specimen the posterior valley is narrow as compared to the C. intermedium, however posterior valley is U-shaped in both the species. Contrary to the present P2, a weak labial cingulum is present in P2 of the C. intermedium. Anterior cingulum is present in the C. intermedium, which is absent in the present collection, and the posterior cingula are present but very much reduced in both the species. Anterior cingulum is present in lower molars of C. intermedium, whereas in the present specimens it is absent. The posterior valley is widely V-shaped lingually whereas it is U-shaped in the present specimens. In M3 trigonid is angularly V-shaped with the 42 narrow and short paralophid and a right angled metalophid which is quite similar to the C. intermedium. No trace of cement is present. There is no labial cingulum. Contrary to the C. intermedium the ectolophid groove is deep and marked to the base of crown in the present collection. If the skull of G. browni is considered in its entirety, and all of its anatomical characters are evaluated, we see that it is seemingly more closely related to the modern Rhinoceros than to any other genera of the Rhinocerotidae. The lower dentition of G. browni follows the general rhinocerotids pattern. PUPC 08/14 and PUPC 08/85 compare favorably with AMNH 29838 (Paratype) (Colbert 1935). They have similar antero- posterior length and crown width and W/L indices (Table 2) with American Museum collection. The specimens morphologically resemble to the species G. browni and metrically the measurements are overlapping the already studied specimens. The slight difference in the measurements is due to the individual variations. It is also clear that measurements lie within the limits of variation. On the basis of above mentioned characteristics it is evident that specimens PUPC 08/14 and PUPC 08/85 belong to genus Gaindatherium. It is, therefore, concluded that on the basis of above mentioned characters like tooth morphology, over all contours of the teeth, size of the teeth, enamel constriction and development of the different crown structures these specimens are referable to the species Gaindatherium browni. Bivariate scatter graphs showing comparison of the studied specimens with the type specimens, Width/length indexes, Size variation of lower dentition are also provided (Plots 3a-c, 4 and 5).

43 Order: ARTIODACTYLA Owen, 1848 Suborder: SUIFORMES Jaeckel, 1911 Superfamily: SUOIDEA (Gray, 1821) Cope, 1887 Family: SUIDAE Gray, 1821 Subfamily: LISTRIODONTINAE Simpson, 1945

Genus : Listriodon Von Meyer, 1846

Type species : Listriodon splendens Von Meyer, 1846

Included species 1. L. splendens Von Meyer, 1846 2. L. lockharti Pomel, 1848 3. L. Intermedius Liu and Lee, 1963 4. L. pentapotamiae Falconer, 1868 5. L. theobaldi Falconer, 1868 6. L. guptai Falconer, 1868 7. L. latidens Biedermann, 1873 8. L. akatikubas Wilkinson, 1976 9. L. juba Ginsburg, 1977 10. L. bartulensis Pickford, 2001 11. L. raetamaensis Pickford and Morales, 2003

Distribution: The genus Listriodon is known from Europe and Africa as well as from the Siwaliks. In Europe it is known in the basal Middle Miocene deposits, in Africa it is known from the Ngorora Formation, and from the Siwaliks it is known from the Chinji formation (Pilgrim, 1926; Pickford, 2001; Pickford and Morales, 2003).

Diagnosis: Lophodont forms of molars are found. Tooth crests are perfect with sharp cutting edges. Teeth are smaller in size than other genera of the family Suidae. Talon in third molar is present and varies in size in different species of the genus and symphysis is also present (Colbert, 1935). 44 Listriodon pentapotamiae Falconer, 1868 Type specimen: GSI B 107, a complete right M2 and fragment of right M3. Also right and left P4.

Locality: Khushalghar below Attock, Punjab (Colbert, 1935).

Stratigraphic Range: Lower Siwaliks and lower portion of the Middle Siwaliks (Colbert, 1935).

Diagnosis: Similar to Listriodon splendens of Europe, but with a larger talon on the third molar, a strong cingulum in the fourth premolar, a shorter and more slender symphysis.

Specimens examined: PUPC 08/9, a right mandibular fragment having P4-M3, a left mandibular fragment having M1-3 and a left side upper incisor; PUPC 08/70, an isolated left lower first molar; PUPC 08/93, an isolated lower left third premolar; PUPC 08/21, an isolated lower right fourth premolar and PUPC 08/24, an isolated right upper canine.

Locality: Dhok Bun Ameer Khatoon, Chakwal district, Punjab, Pakistan.

Stratigraphic range: Lower Siwaliks (Chinji Formation).

Description

PUPC 08/9 Mandibles (Figure 8&9) The mandibles are unworn and were collected from the same place, suggesting that they came from young animals, and possibly the same individual. They are very well preserved and both ramii are thick transversely and deep vertically. The anterior and posterior parts of the right mandible (Figure 8) are missing preserving P4-M3. Total molar length series of right mandible is 78 mm. The depth below

M2 is 32 and the width below M2 is19 mm. Preserved horizontal ramus is 89.7 mm.

P4 is well preserved with quite thin and shiny enamel. Since the tooth is unworn, nothing can be said about the thickness of the enamel and dentine. It is low crowned with pointed conids. There are only two cusps on the anterior side arranged in a transverse line. The inner cusps are much stronger. Strongly developed anterior and posterior 45 cingula are present. The anterior cingulum is much stronger than posterior one. M1 is finely preserved and the enamel shows a uniform thickness all over the crown of tooth.

Little or no wearing is observed in M2. The enamel is smooth and not rugose. M3 is badly damaged anteriorly. The protoconid and metaconid are broken and only base is present. The tip of the hypoconid is also broken. The sharp chisel shaped crests are present. These crests are perfect with very sharp cutting edges. A large talonid is present and is of equal height to the other conids of the molar. The protoconid is attached to the metaconid and a loph is formed. Similarly a loph between the hypoconid and entoconid is also formed. At the anterior side of the protoloph a depression is present. The protoconid and hypoconid and also the metaconid and entoconid are separated by a deep valley. There is a downward slope from the hypoconid to the metaconid. All four conids of the molars are pointed at the top. In general appearance all the conids are V shaped. The protoconid is vertically higher than the metaconid. No conules are visible. The metaconid is high compared to other conids. The protoconid and hypoconid are equal in height. The entoconid is less high than the metaconid but higher than the protoconid and hypoconid. On the anterior side of the molars a deep depression is present and it makes a V shape valley in the mid and anterior side of the protoloph.

Left mandible (Figure 9) consists of M1-3 and is also broken anteriorly. The remaining parts are very well preserved. It is moderately thick transversely and deep vertically. The total length of the molar series is 61 mm. The depth of the mandible below the M2 is 32 mm and the width is 19 mm, and M3 is 22 mm. The preserved horizontal ramus is 63 mm. The preserved ascending ramus is 22 mm and the reconstructed ascending ramus is 37 mm. The protoconid is joined with the metaconid, and the hypoconid is joined with the entoconid, forming lophs. All the molar series is well preserved. The wear is more confined to the protoconid and hypoconid as compared to metaconid and entoconid in M1. Metaconid and entoconid are looking high than other conids. The apex of metaconid is slightly damaged in M2. M3 is well preserved and unworn. Smooth, single layered, shinny and thin enamel is present. In M3 large talonid is present posteriorly and all the four conids and talonid are pointed at the top with sharp chisel shaped crests. On the anterior side of tooth and around the talonid and on the buccal side between the protoconid and hypoconid cingulum is present. Anterior and posterior cingulum is well developed. 46

PUPC 08/93 (Figure 10) It is an isolated lower left third premolar. It is well preserved with quite thin and shinny enamel. The tooth is worned, so the thickness of enamel is 1.5 mm. The enamel is not wrinkled or rugose. The tooth is in the middle stage of wear. According to measurements (Table 3) the tooth is low crowned with pointed conids. There are only two cusps on the anterior side arranged in a transverse line. The inner cusp is very strong as compared to outer one. Anterior and posterior cingulum is strongly developed. The anterior cingulum is much stronger than posterior one. PUPC 08/21(Figure 11) It is an isolated fourth lower right premolar well preserved with quite thin and shinny enamel. Since the tooth is unworn, so nothing could be said about the thickness of enamel and dentine. The enamel is not wrinkled or rugose. The tooth is in an early stage of wear. Tooth is low crowned with pointed conids having only two cusps on the anterior side arranged in a transverse line. Tooth has strongly developed anterior and posterior cingulum. PUPC 08/70 (Figure 12) It is an isolated left lower first molar which is in the middle stage of wear. It shows that the enamel is thin, shinny all around the crown. PUPC 08/24 (Figure 13) It is an isolated right upper canine. It is almost complete anteriorly and missing posteriorly. The antero-posterior preserved length is 30 mm and transverse width is 17 mm (Table 3). It is entirely composed of dentine and probably is in the middle stage of wear. The length of wear stage from anterior to posterior is 23 mm and width is 16 mm. It is in an excellent stage of preservation. The cross section of the canine from the posterior side is circular as shown by diagram. In the center of enamel ridge from anterior to posterior is prominent and it is covered laterlly by thin layer of bone. PUPC 08/09 (Figure 14) The left upper incisor I1 is chisel shaped, it is well preserved and the enamel is shiny. It is embedded in the jawbone. The single cusp is sharp and eroded. It is in an early stage of wear and its physical characteristics also confirm this status. Its crown length is 18 mm, width 8.2 mm and height 11 mm. A small portion of bone in which incisor is inserted i.e. the premaxilla is well preserved. Its length is 37.5 mm and its width is 19.5 mm. Symphysis length is 19 mm and width 7 mm.

47

A

B

C

Figure 8: L. pentapotamiae, PUPC 08/9, right mandibular fragment having P4-M3, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

48

A

B

C

Figure 9: L. pentapotamiae, PUPC 08/9, left mandibular fragment having M1-3, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

49

A

B

Figure 10: L. pentapotamiae, PUPC 08/93, left third lower premolar, A - occlusal view, B - anterior view. Scale bar 10 mm.

50

A

B

C

Figure 11: L. pentapotamiae, PUPC 08/21, right fourth lower premolar, A- lingual view, B - occlusal view, C - posterior view. Scale bar 10 mm. 51

Figure 12: L. pentapotamiae, PUPC 08/70, left first lower molar. Scale bar 10 mm.

A

B

Figure 13: L. pentapotamiae, PUPC 08/24, right upper canine, A - buccal view, B - lingual view. Scale bar 10 mm. 52

A

B

Figure 14: L. pentapotamiae, PUPC 08/9, left upper incisor, A - occlusal view, B - lingual view. Scale bar 10 mm. 53 Table 3: Measurements of teeth referred to L. pentapotamiae Falconer, 1868. (Data compared with American Museum specimens). *The studied specimens.

Number Position Length Width W/L

PUPC 08/9* I1 18 8.1 0.45

PUPC 08/24* C 30 17 0.57

PUPC 08/93* P3 13 11.5 0.88

PUPC 08/21* P4 15 11 0.73

PUPC 08/9* P4 15.2 10.6 0.70

PUPC 08/9* M1 16 13 0.81

PUPC 08/70* M1 14 13 0.93

AMNH 19519 M1 16 12 0.75

AMNH 19457 M1 17 13.5 0.79

AMNH 19929 M1 16 12 0.75

PUPC 08/09* M2 22 16 0.73

AMNH 19519 M2 19 16 0.84

AMNH 19624 M2 23 18 0.78

AMNH 19625 M2 19.5 14.5 0.74

AMNH 19629 M2 18 14 0.78

AMNH 19432 M2 21 15 0.71

AMNH 19541 M2 19 13 0.68

PUPC 08/9* M3 28 16 0.57

AMNH 19432 M3 28 16 0.57

AMNH 19519 M3 27.5 18 0.65

AMNH 19542 M3 28.5 17 0.60

AMNH 19625 M3 28 16 0.57

54

Lower P4

11.05 11 10.95 10.9 10.85 PUPC 08/21* 10.8 PUPC 08/9* Length 10.75 10.7 10.65 10.6 10.55 14.95 15 15.05 15.1 15.15 15.2 15.25 Width

a

Lower M1

13.6 13.4 13.2 PUPC 08/9* 13 PUPC 08/70* 12.8 AMNH 19519 12.6 Length AMNH 19457 12.4 AMNH 19929 12.2 12 11.8 0 5 10 15 20 Width

b

Lower M2

20 18 16 PUPC 08/09* 14 AMNH 19519 12 AMNH 19624 10 AMNH 19625

Length 8 AMNH 19629 6 AMNH 19432 4 AMNH 19541 2 0 0 5 10 15 20 25 Width

c

Lower M3

18.5

18

PUPC 08/9* 17.5 AMNH 19432 17 AMNH 19519

Length AMNH 19542 16.5 AMNH 19625 16

15.5 27.4 27.6 27.8 28 28.2 28.4 28.6 Width

d Plot 6 a-d: Bivariate scatter graphs showing comparison of the studied specimens with the type specimens. 55

20

18

16

14 PUPC 08/9* (I1) PUPC 08/24* (C**) PUPC 08/93* (P3) PUPC 08/21* (P4) 12 PUPC 08/9* (P4) PUPC 08/9* (M1) PUPC 08/70* (M1) AMNH 19519 (M1) 10 AMNH 19457 (M1) AMNH 19929 (M1)

PUPC 08/09* (M2) AMNH 19519 (M2) Length 8 AMNH 19624 (M2) AMNH 19625 (M2) AMNH 19629 (M2) AMNH 19432 (M2) 6 AMNH 19541 (M2) PUPC 08/9* (M3) AMNH 19432 (M3) AMNH 19519 (M3) 4 AMNH 19542 (M3) AMNH 19625 (M3)

2

0 0 5 10 15 20 25 30 35 Width

Plot 7: Shows the size variation of cheek teeth L. pentapotamiae

W/L index 1

0.9

0.8

0.7 I1 0.6 C** P3 0.5 P4

0.4 M1 M2 0.3 M3

0.2

0.1

0 1 2 3 4 5 6 7

Plot 8: Width/length ratio of studied specimens L. pentapotamiae 56 Table 4: Dental measurements of different species of the genus Listriodon (in mm) taken from Picford and Morales, (2003) and Orliac et al. (2009).

Species Number Position Length Width W/L L. guptai PAK 2269 I1 19 11.3 0.59

L. guptai PAK 1389 P3 16.3 10.7 0.66

L. guptai PAK 1386 P4 16.8 11.5 0.68

L. guptai PAK 1385 P4 17.2 13 0.76

L. guptai PAK 1385 M1 19.2 13.3 0.69

L. guptai PAK 1385 M2 21.8 16.7 0.77

L. guptai PAK 1385 M3 29.2 28.3 0.97

L. retamaensis RET 17 P4 16.5 11.4 0.69

L. retamaensis RET 2 M1 14e 13.9e 0.99

L. retamaensis RET 2 M2 19.0 15.9 0.84

L. retamaensis RET 2 M3 31.5 17.9 0.57

L. retamaensis AC 163 P3 12.6 8.0 0.63

L. retamaensis AC 163 P4 15.6 11.0 0.71

L. retamaensis AC 163 M3 29.6 17.3 0.58

L. retamaensis EO 180 P3 14.5 8.9 0.61

L. retamaensis VD 61 M3 34.0 17.4 0.51

L. locharti _ P3 18.4 10.5 0.57

L. locharti _ P3 18.7 10.5 0.56

L. locharti _ P4 17.8 12.6 0.71

L. locharti _ P4 19.0 13.1 0.69

L. locharti _ M1 20.4 14.2 0.70

L. locharti _ M1 20.4 15.7 0.77

L. locharti _ M2 23.4 19.0 0.81

L. locharti _ M2 23.6 18.6 0.79

L. locharti _ M3 30.5 19.0 0.62

L. locharti _ M3 31.0 18.6 0.6

L. splendens _ P3 17.0 9.6 0.56

L. splendens _ P3 15.9 10.1 0.64

L. splendens _ P4 16.4 13.3 0.81

L. splendens _ P4 16.7 12.9 0.77

L. splendens _ M1 16.4 12.5 0.76

L. splendens _ M2 20.8 16.3 0.78

L. splendens _ M3 31.9 19.6 0.61 57

Lower I1

12

10 L. pentapotamiae PUPC 08/9*

L. guptai 8

6 Length

4

2

0 17.8 18 18.2 18.4 18.6 18.8 19 19.2 Width a

Lower P3

14

12 PUPC 08/93* L. guptai

10 L. retamaensis L. retamaensis

L. locharti L. locharti 8 L. splendens L. splendens

Length 6

4

2

0 0 5 10 15 20 Width

Lower P4

14

12 PUPC 08/21* L. guptai

10 L. guptai L. retamaensis L. retamaensis L. locharti

8 L. locharti L. splendens

L. splendens

Length 6

4

2

0 0 5 10 15 20 Width b 58

Lower M1

18

16

PUPC 08/9* PUPC 08/70* 14 AMNH 19519 AMNH 19457

12 AMNH 19929 L. guptai

10 L. retamaensis L. locharti L. locharti L. splendens

Length 8

6

4

2

0 0 5 10 15 20 25 Width c

Lower M2

20

18

16 PUPC 08/09* AMNH 19519 AMNH 19624 AMNH 19625 14 AMNH 19629 AMNH 19432

12 AMNH 19541 L. retamaensis L. locharti L. locharti 10

L. splendens Length 8

6

4

2

0 0 5 10 15 20 25 Width d

Lower M3

30

25 PUPC 08/9* AMNH 19432

AMNH 19519 AMNH 19542 20 AMNH 19625 L. guptai

L. retamaensis L. retamaensis L. retamaensis L. locharti 15

L. locharti L. splendens Length

10

5

0 0 5 10 15 20 25 30 35 40 Width e

Plot 9a-e: Variation in the dentition of various species of Listriodon. 59 Discussion

The sharp chisel shaped crest is a major feature of the molar teeth of Deinotheriids, proboscideans, lophodonts pigs and some metatheres. The teeth under discussion are too small to be referred to any of the proboscideans. In lophodonts metatheres the crest is imperfect. In lophodont pigs i.e. listriodonts, the tooth crests are perfect with very sharp cutting edges. The specimen (PUPC 08/9) consists of incisor, premolar and molars inserted into right and left mandibuler ramii. They probably belong to same individual. The specimens under study comprise favorably with Ind. Mus. B 697 and AMNH 19519 described by Pilgrim (1926) and Colbert (1935). The teeth are in an early wear stage. Protoconid inferred to have been largest cusps. Metaconid located distally in respect to protoconid but does not invade labial basin. Under the heading of Listriodon pentapotamiae Colbert (1935) described the type 2 3 specimen GSI B107, a complete right M and fragment of right M , also right and left P4.

He described AMNH 19457, AMNH 19928 having M1 and AMNH 19541, AMNH

19624, AMNH 19629, having M2 while AMNH 19519 having M1-3. These all specimens were collected from Phadial, Rammagar and Nathot and near the Chinji rest house area. Colbert described these specimens as L. pentapotamiae. Specimens PUPC 08/9, PUPC 08/21, PUPC 08/24, PUPC 08/70 and PUPC 08/93 are collected from Dhok Bun Ameer Khatoon, District Chakwal, Chinji formation (Lower Siwaliks). After thorough investigation and measurements of studied specimen with AMNH 19519, it is evident that both specimens have same features and slight differences in size can only be attributed to individual variation. The material under study (PUPC 08/9) consists of right and left mandibular ramii of young individual as shown by teeth wear. Actually preserved anterior posterior horizontal ramus in right side mandible is 89.7 mm. It is moderately thick transversely and deep vertically. Width of mandible below the M2 is 19 mm and M3 is 22 mm. Preserved horizontal ramus is 63 mm. Preserved ascending ramus is 22 mm and reconstructed ascending ramus is 37 mm. Colbert (1935) did not mention this feature. The teeth under study are lophodont because lophs are formed between protoconid and metaconid, hypoconid and entoconid. In this respect the specimens under study resembles with Amer. Mus. collection. They show all the basic features of the species i.e. prominent lophs, M3 having two cross crests followed by talonid. A faint cingulum is present on the anterior side. This cingulum and the longitudinal ridges are also seen in the diagram by Pilgrim (1926), though not mentioned by him. 60

M1 and M2 are similar to those of Ind. Mus. B. 697 and AMNH 19519 referred by Pilgrim (1926) and Colbert (1935) respectively. Both of these teeth have two transverse ridges connected by a diagonal ridge. Prominent cingula are present on the anterior and posterior side. Tooth with crests having sharp edges. A strong cingulum in fourth premolar is present. These entire features are similar to those of Ind. Mus. B 697.

However, the diagonal ridge is not visible in PUPC 08/9. PUPC 08/93 is an isolated P3 while right mandibular P4 and an isolated P4 (PUPC 08/21) have only two cusp on the anterior side arranged in a transverse line. These are followed by a low talonid. This structure is similar to the specimen described by Pilgrim (1926) and Colbert (1935). Among early workers Colbert, 1935 described a specimen AMNH 19586 as I1. Left upper incisor has same structure as Colbert (1935) described in figure i.e. Colbert p.233 in AMNH 19586. This compares favorably with incisor (PUPC 08/9) in size and general appearance. Pilgrim (1926) mentioned a specimen as I1, P-32, but does not refer it to any species. Since the specimen PUPC 08/9 is very well preserved and is showing the morphological features clearly. Molars of the both mandibular ramii also resemble each other in all the structural details and in antero-posterior length and crown width as given in Table 3 & 4. On the basis of these similarities and resemblances the specimens under study are being referred to the species L. pentapotamiae Falconer, 1868. Dental measurements of different species of the genus Listriodon taken from Picford and Morales, (2003) and Orliac et al. (2009) are also given in table 4, which indicates that teeth of L. pentapotamiae are smaller in size comparatively. Plots (6a-d, 7 and 8) are also provided having width/length ratio of studied specimens, comparison of the studied specimens with the type specimens and the size variation of cheek teeth of L. pentapotamiae while Plot (9a-e) indicate variation in the dentition of various species of Listriodon.

61 Suborder: RUMINANTIA Scopoli, 1777 Family: TRAGULIDAE Milne-Edwards, 1864

Genus: Dorcatherium Kaup, 1833

Type Species: Dorcatherium naui Kaup, 1833

Included Species 1. D. naui Kaup, 1833 2. D. vindobonense Von Meyer, 1846 3. D. guntianum Von Meyer, 1846 4. D. crassum Lartet, 1851 5. D. minus Lydekker, 1876 6. D. majus Lydekker, 1876 7. D. jourdani Deperet, 1887 8. D. peneckei Hoffman, 1893 9. D. birmanicus Noetling, 1901 10. D. puyhauberti Arambourg and Piveteau, 1929 11. D. chappuis Arambourg, 1933 12. D. songhorensis Whitworth, 1958 13. D. parvum Whitworth, 1958 14. D. pigotti Whitworth, 1958 15. D. bulgaricum Bakalov and Nikolov, 1962 16. D. nagrii Paresad, 1970 17. D. libiensis Hamilton, 1973 18. D. minimus West, 1980 19. D. orientale Qiu Zhanxiang and Gu Yumin, 1991 20. D. moruorotensis Pickford, 2001 21. D. iririensis Pickford, 2002 62 Distribution: The genus Dorcatherium is known from the Lower Miocene of Europe by Kaup (1833) and Arambourg and Piveteau (1929). It is also reported from the Miocene deposits of East Africa by Lartet (1837), Arambourg (1933), Whitworth (1958) and Hamilton (1973). It was first reported from the Siwalik Hill of Pakistan by Lydekker (1876) and has since been reported by numerous authors including Colbert (1935), Sahni et al. (1981) West (1980) Farooq (2006) and Farooq et al. (2007a,b &2008). I have described specimens from Dhok Bun Ameer Khatoon, Chakwal District. Diagnosis: The upper molars have strongly developed buccal styles and a basal cingulum. The lower molars are characterized by a well developed or vestigial median basal pillar and a well developed or vestigial ectostylid. A dorcatherium fold is also present (Khan et al., 2005).

Dorcatherium majus Lydekker, 1876

Type Specimen: GSI B 195, two upper molars namely right M1-2.

Locality: Dhok Bun Ameer Khatoon, Chakwal district, Punjab, Pakistan.

Stratigraphic range: Lower Siwaliks (Chinji Formation).

Locality: Khushalgar, near Attock. Also Hasnot and Adjacent Localities (Colbert, 1935)

Stratigraphic range: Lower and Middle Siwaliks.

Diagnosis: Dorcatherium majus is equal in size to Dorcabune anthracotherioides but larger than D. minus. It is characterized by a strong mesostyle and parastyle; the upper molars have a well-developed cingulum and are extremely hypsodont. In the lower molars a stoutly developed ectostylid and well developed median basal pillar are present (Farooq et al., 2008).

Distribution: Dhok Bun Ameer Khatoon, district Chakwal, Punjab, Pakistan.

Specimens examined: PUPC 08/23, right maxillary fragment having M2-3; PUPC 08/42, right second upper molar; PUPC 08/36, an isolated second right upper molar and PUPC 08/39, right lower first lower molar. 63 Description Upper dentition M2 PUPC 08/36 (Figure 16) consists of an isolated right upper molar. It is moderately well preserved. Its buccal and posterior sides are broken. It is embedded in bone whose length is 23 mm and width is 22 mm. PUPC 08/42 (Figure 17) is an isolated left second upper molar. It is very well preserved. In general appearance both teeth look quadrate and are transversely wider anteriorly than posteriorly (Table 5). Regarding the antero- posterior length, the specimens are more elongated towards the lingual side than the buccal side. The molars are hypsodont with narrow crowns as indicated by the W/L indices. The enamel is wrinkled and uniformly thick with a thickness of 1.0 mm. The cingulum is weakly developed anteriorly and strongly developed posteriorly in PUPC 08/42 while in PUPC 08/36 the posterior side is damaged and the cingulum cannot be observed. It is also present on the inner side especially where it covers the base of the protocone. All four cusps are inclined towards the median longitudinal axis of the molar. This inclination is more prominent in the inner cones than the outer cones. The protocone is almost L-shaped, and its posterior end does not join at the middle of the hypocone. The inner most lingual part of the protocone is rounded. The protocone is semi-crescentic in shape and its anterior arm is longer than the posterior arm. Its anterior arm is joined with the parastyle by a thin ridge of enamel while the posterior arm is free. The paracone is relatively smaller in antero-posterior length compared to the metacone. The parastyle and mesostyle cannot be observed in PUPC 08/36 because its buccal and posterior sides are broken. The metacone is the highest among all the cones. It is broken anteriorly and posteriorly in PUPC 08/36 while in PUPC 08/42 it is very well preserved. The posterior half of the paracone is thicker than the anterior half. The parastyle is well developed and connected to the base of the anterior rib. The hypocone is more crescentic and both of its outer ends are narrow. It is less worn than the protocone. Both of its arms are equal in length, exhibiting a V-shaped structure. Its anterior arm is joined with the posterior half of the paracone while its posterior arm remains free. PUPC 08/23 (Figure 15) is an excellent part of the right maxilla bearing M2-3. The whole part of the maxilla is broken away in front of the teeth. The antero-posterior length of the teeth is 32 mm. The jaw length in which M2-3 is embedded is 35 mm and the width is 19 mm. The molars are finely preserved and are almost half worn. The teeth are squarish in general appearance. 64 M3 The molar is well preserved and is almost half worn. The enamel is rugose at all sides and its average thickness is 1.1 mm. A strong horizontal fold forms a cingulum at the lingual side. The cingulum is moderately developed at the anterior and posterior face of the molar but is comparatively well developed towards the inner side, especially at the entrance of the transverse valley between the protocone and hypocone. The central cavities are deep and wide. The molar is brachydont and narrow crowned as shown by measurements in Table 5. The protocone is more worn than all the other cones and is almost L-shaped. Its posterior end does not join at the middle of the hypocone. Its anterior arm is longer than the posterior arm and anterior arm joined with the parastyle by a thin ridge of enamel while the posterior arm is free. The inner most lingual part of protocone is blunt and is almost rounded at this stage of wear. The paracone is comparatively smaller than metacone in antero-posterior length. The parastyle is very well developed. It is not vertical but inclined. The median rib is stoutly developed. The mesostyle is well developed and divergent. The metacone is very thin on its upper end and becomes thick towards its base. Its anterior and posterior arms are inclined towards the labial side. Its median ribs are moderately developed. It is not vertical but slightly inclined towards the anterior rib. All four cusps are inclined towards the median longitudinal axis of the molar, although the degree of inclination is greater in the lingual cusps than the buccal cusps. The metacone is roughly concave at the outer side. The metastyle is weakly developed. The hypocone is more crescentic and both of its outer ends are narrow. It is less worn than the protocone. Both of its arms are equal in length, giving a V-shaped structure. Its anterior arm is joined with the posterior half of the paracone while its posterior arm remains free. The median ribs are well developed with the anterior rib being the strongest among the styles.

Lower dentition

M1 PUPC 08/39 (Figure 18) is a very well preserved first right lower molar. The maximum preserved height of bone under the M1 is approximately 14 mm and the thickness is about 9 mm. The antero-posterior length of bone below M1 is 19 mm. It is in an early stage of wear. The enamel is moderately thick and rugose. The protoconid is more worn than the other conids. The ectostylid is moderately developed and seems to be 65 associated with the posterior side of the protoconid. The buccal and lingual sides of the molar show very fine placations and an almost uniform thickness all over the crown, with an average thickness of 1.0 mm. The central cavities are narrow antero-posteriorly. The ectostylid is very small and unworn exhibiting a close association with the postprotocristid. The protoconid is antero-posteriorly longer than the metaconid. It is crescentic in shape with a preprotocristid and a postprotocristid. The postprotocristid is bifurcated at the point where it is attached to the postmetacristid. The dentinal islet of the protoconid is roughly triangular. The dorcatherium fold is present in the molar. The metaconid and the entoconid are higher than their opposite cusp. The metaconid is pointed in the middle with anterior and posterior sloping ridges. The postmetacristid is also slightly bifurcated by a vertical groove and is slightly more worn than the premetacristid. The premetacristid has produced a very narrow but elongated dentinal islet. The entoconid is well developed. The hypoconid is more crescentic than the protoconid. The hypoconulid is very well developed and lies in line with the lingual conids. It is crescentic in appearance. Its median valley is shallow and opens towards the buccal side. The cingulid is lacking lingually, whereas antero-posteriorly it is strongly developed and high. The conids are well developed and pointed at the apex, and the dorcatherium fold is present in the postprotocristid. The degree of wear is greater in the buccal conids than in the lingual conids. The buccal conids are selenodont, whereas the lingual ones have a trend towards the conical form. The postprotocristid is deeply bifurcated into two daughter cristids, of which the outer one is free and the inner one is linked with the postmetacristid. 66

A

B

C

Figure 15: D. majus, PUPC 08/23, right maxillary fragment having M2-3, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

67

A

B

C

Figure 16: D. majus, PUPC 08/36, an isolated right second upper molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

68

A

B

C

Figure 17: D. majus, PUPC 08/42, an isolated right second upper molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

69

A

B

C

Figure 18: D. majus, PUPC 08/39, right first lower molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

70 Table 5: Comparative Measurements (in mm) of molar teeth referred to D. majus Lydekker, 1876. *The studied specimens (AMNH taken from Colbert, 1935).

Number Position Length Width W/L

PUPC 08/23* M2 16 17 1.06

PUPC 08/36* M2 16 14 0.87

PUPC 08/42* M2 15.5 13 0.84

PUPC 08/23* M3 17 17.5 1.04

AMNH 19302 M2 18.5 21.5 1.42

AMNH 19354 M3 20.5 23.5 1.15

GSI B 198 M2 19.6 19.6 1.0

GSI B 198 M3 20.1 19.2 0.96

PUPC 67/191 M2 13.3 14.5 1.09

PUPC 68/33 M2 13.3 14.5 1.09

PUPC 68/250 M2 15.7 16.5 1.05

PUPC 85/15 M2 19 20 1.05

PUPC 85/21 M2 18 22 1.22

PUPC 87/328 M2 17.7 19 1.07

AMNH 19302 M2 18.5 21.5 1.16

PUPC 67/191 M3 13.65 15.25 1.12

PUPC 87/197 M3 20.5 22 1.07

PUPC 87/328 M3 19.1 18.2 0.95

PUPC 08/39* M1 13 8 0.62

AMNH 19520 M1 14 9 0.64

AMNH 19524 M1 13.5 9 0.67

GSI B M1 15.7 9.5 0.61

PUPC 86/2 M1 14.3 9 0.63

PUPC 86/5 M1 13 9.3 0.72 71

Upper M1

9.6 9.4

9.2 PUPC 08/39* 9 AMNH 19520 8.8 AMNH 19524

8.6 GSI B Length 8.4 PUPC 86/2 8.2 PUPC 86/5 8 7.8 0 5 10 15 20 Width

a

Upper M2

25 PUPC 08/23* PUPC 08/36* 20 PUPC 08/42* AMNH 19302 15 GSI B 198 PUPC 67/191 Length 10 PUPC 68/33 PUPC 68/250 5 PUPC 85/15 PUPC 85/21 0 0 5 10 15 20 25 PUPC 87/328 AMNH 19302 Width

b

Upper M3

25

20 PUPC 67/191 PUPC 87/197 15 PUPC 87/328 GSI B 198 Length 10 AMNH 19354 PUPC 08/23* 5

0 0 5 10 15 20 25 Width

c Plot 10a-c: Bivariate scatter graphs showing comparison of the studied specimens with the type specimens. 72

W/L index

1.6

1.4

1.2

1 M2 0.8 M3 M1 0.6

0.4

0.2

0 1 2 3 4 5 6 7 8 9 10 11 12

Plot 11: Width/length indexes of the cheek teeth D. majus.

73 Discussion

In larger species of Dorcatherium, such as D. peneckei and D. vindobonense, the cheek teeth are more bunodont. Relative to their breadth, premolars are usually longer than the recent tragulids. The upper canines of males are large and trenchant (Whitworth, 1958). At least a vestige of an ectostylid is present in almost all forms of the genus; in some, the ectostylid is even rather robust and very prominent. The external styles and the basal cingulum are well developed in the upper molars (Colbert, 1935). Dorcatherium resembles the extant African genus Hyaemoschus in terms of the presence of a first lower premolar but it is often absent in D. crassum (Whitworth, 1958). Dorcatherium also has less selenodont cheek teeth, a cingulum, more robust jaws, and a contact between the premaxilla and nasals. Before the Eurasian and African Miocene Dorcatherium the evolutionary history of tragulids was unknown (Webb and Taylor, 1980). Dorcatherium is the first undoubted fossil tragulid from Africa and Eurasia. The numerous species referred to Dorcatherium mainly differ by their size (West, 1980). D. chappuisi differs from D. minimus from Pakistan by the presence of a cingulum on the anterior and lingual sides of the upper molars and due to its slightly smaller size. Paracone and metacone of D. minimus have greater buccal flare than in D. moruorotensis. Indeed where D. minimus differs from D. moruorotensis, it approaches the genus Tragulus (West 1980). Whitworth (1958) provided measurements of several specimens of D. parvum from Rusinga Island, Kenya (17.8 Ma), Upper third molars of D. minimus from Daud Khel, Pakistan are slightly larger than those of D. moruorotensis. The Daud Khel molars have no cingulum on the protocone, unlike the condition in D. moruorotensis, and they may belong to Tragulus rather than Dorcatherium (West, 1980). D. naui, first described from Eppelsheim, Germany in that its lingual cusps are more fully crescentic, the outer walls more nearly vertical, the labial rib of the metacone reduced, the enamel thinner, the mesostyle more prominent, the parastyle projecting less forwards, and the cingula weaker and smoother surfaced (Fahlbusch, 1985). There are three species of the genus Dorcatherium in the Siwalik Hills of Pakistan: i) - D. majus ii) - D. minus iii) - D. minimus. Both D. majus and D. minus show the same characteristics but vary in their size, with D. majus being larger than D. minus. In the genus Dorcatherium the protocone is an incomplete crescent and the median ribs of the paracone are forwardly directed producing a notch anterior to it. The premolar row is comparatively larger than the molar row. Compared to modern tragulids the molar teeth in the large Dorcatherium species are more bunoid, and the premolars are usually 74 longer relative to their breadth (Whitworth, 1958). Dorcatherium seems to be the genus of the family Tragulidae that has developed analogously to the recent African genus Hyaemoschus. The basal cingulum is well developed in Dorcatherium, and at least a vestige of an ectostylid remains in the lower molars. The specimens PUPC 8/23, PUPC 08/42 and PUPC 08/36 bearing second right and left molars are smaller in size with regards to width and length compared to those of the type specimen (AMNH 19302), although their indices of W/L ratio are very close to the type specimen. Second upper molars comprise both worn and unworn teeth. Morphological characters and dental measurements have been studied and checked for a resemblance with the type specimen in the structure of the cusps, development of styles, cingula and antero-posterior length. The differences in these dental measurements are quite insignificant because their indices show that all specimens resemble each other. A common feature of these teeth is the development of a mesostyle as mentioned by Colbert (1935), which is a very strong isolated pillar. M3 measurements exhibit a close relationship with the type specimen AMNH 19354 (Colbert, 1935) and GSI B 198 (Pilgrim, 1914). The upper last molar resembles the type specimen in the structure of the cusps, cingula, rugosity of enamel, antero-posterior length and the development of styles. The specimen is different in its length and width from both AMNH 19354 and GSI B 198, but the W/L index lies close to specimen AMNH 19354 (i.e. both are broad crowned). The specimens under study show all the morphological features of the cheek teeth cited by Lydekker (1876) and Colbert (1935). The upper molars are distinguished by their squarish appearance and their stronger development of parastyles, mesostyles and cingula. The metastyle is always weakly developed. The lingual cusps seem to be more inclined towards the median longitudinal axis than the buccal ones. The enamel is plicated and the central cavities are well developed. All these characteristics are exhibited by the present collection of the upper dentitions (PUPC 08/23, PUPC 08/42 and PUPC 08/36) indicating that the studied specimens belong to D. majus. The comparisons of the dental measurements of the specimens under study with those of the type specimens also show a resemblance and very slight differences. The molars compare very closely to those of specimens AMNH 19302, AMNH 19354 (Colbert 1935) and GSI B 198 (Pilgrim, 1915) (Table 5). They are therefore referred to the genus Dorcatherium and species D. majus. The lower molars of D. majus are comparatively high and narrow crowned (Colbert 1935). They have ectostylids between the protoconids and hypoconids. The 75 metaconid is recognized as the highest one among all the conids. Stylids are weakly or improperly developed. Their anterior median ribs are well pronounced. All these characteristics exhibited by the present specimen of the lower dentition indicate very clearly that it belongs to the species D. majus (Colbert, 1935). PUPC 08/39 seems to have a close resemblance regarding its length and width to specimens AMNH 19524 (Colbert, 1935) of the type material and AMNH 19520, but differs markedly in length from the specimen GSI B 593 of the type material (Table 5). In spite of this difference in length, the W/L indices of all the specimens under study and the type specimen demonstrate that they all are narrow crowned and the differences in length are in a range of intraspecific variability. Their measurements with respect to length and width indices compare well with those of specimens AMNH 19520, AMNH 19524 (Colbert, 1935), and GSI B 593 (Pilgrim 1910) (Plots 10a-c and 11). 76 Dorcatherium cf. minus Lydekker 1876

Type Specimen: Right M1-2 (GSI B 195).

Locality: Near Hasnot, Jhelum district, Punjab, Pakistan.

Stratigraphic range: Lower and Middle Siwaliks.

Diagnoses: A small species of the genus Dorcatherium with sub-hypsodont and broad crowned molars having well developed cingulum, rugosity, styles, moderately developed ribs and vestigial ectostylids.

Known Distribution: Chinji, district Chakwal; Nathot and Phadial, district Jhelum, Punjab, Pakistan; Dhulian, district Attock, Punjab, Pakistan. (Khan et al., 2005) and Dhok Bun Ameer Khatoon, district Chakwal, Punjab, Pakistan by the present author.

Specimen Examined: PUPC 08/90, an isolated first left upper molar.

Locality: Dhok Bun Ameer Khatoon, Chakwal district, Punjab, Pakistan.

Stratigraphic range: Lower Siwaliks (Chinji Formation).

Description

PUPC 08/90 (Figure 19) It is an isolated first upper left molar and is very well preserved. It is subhypsodont and broad crowned (Table 6). In general appearance, it is quadirate and transversely wider anteriorly than posteriorly. The enamel is 0.4 mm thick and very nicely plicated all around the specimen. The cingulum is well developed around the protocone. The outer cones are higher vertically than the inner ones. The protocone situated anteriorly, lingually and more worn as compared to all other cones. It is almost L- shaped and less crescentic because its inner border is rounded and outer one is straight. Its anterior arm is longer than the posterior one and joined with parastyle with the help of a thin ridge of enamel while posterior arm is free and does not join at the middle of 77 hypocone. The parastyle is well developed and connected to the base of the anterior rib. The mesostyle is well developed and is more associated with the metacone. Hypocone is more crescentic and the anterior side is slightly pinched due to pressure of the posterior end of the protocone. It is less worn than the protocone. Both of its arms are equal in length, exhibiting V-shaped structure. Anterior median rib is stronger as compared to posterior one. 78

A

B

C

Figure 19: D. cf. minus, PUPC 08/90, left first upper molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

79

Table 6: Comparative Measurements (in mm) of left M1 referred to D. cf. minus Lydekker, 1876. *Studied specimens (AMNH taken from Colbert, 1935).

Number Position Length Width W/L

PUPC 08/90* M1 9.4 10 1.06

AMNH 19517 M1 12 11 0.92

AMNH 29856 M1 9.8 10 1.0

GSI B 195 M1 10 10 1

PUPC 68/355 M1 9.2 10.2 1.12

PUPC 87/40 M1 10 11.7 1.17

PUPC 87/84 M1 93 10 0.11

PUPC 95/01 M1 9.3 9 0.97

PUPC 02/01 M1 8 10 1.25

PUPC 04/3 M1 9.6 6.1 0.64

Upper M1 14 PUPC 08/90* 12 AMNH 19517 AMNH 29856 10 GSI B 195 8 PUPC 68/355 PUPC 87/40 6 Length PUPC 87/84 4 PUPC 95/01 PUPC 02/01 2 PUPC 04/3 0 0 20 40 60 80 100 Width

Plot 12: Bivariate scatter graph showing comparison of the studied specimens with the type specimens and previously studied specimens.

80

Discussion

The genus Dorcatherium was named by Kaup (1833). Dorcatherium naui Kaup is the type species and is known from Late Miocene deposits of Eppelsheim in Germany (Kaup 1833, Gentry 1978). Dorcatherium is one of the 5 extinct genera of the family Tragulidae along with Dorcabune, Siamotragulus, Yunnanotherium, and Archaeotragulus (Farooq et al., 2008). The dental formula of the genus Dorcatherium is 0.1.3.3/3.1.4.3. The distinct features of the cheek teeth given by Kaup (1833) and elaborated by Zittel (1964) are that the upper molars are quadritubercular and the lower molars have ridges on the metaconid and the protoconid. Piveteau (1961) gave a more detailed generic description, in which the upper molars are quadrangular in form, a well–developed cingulum surrounds the protocone, cusps are conical and the teeth tend to be hypsodont. The first lower premolar is sometimes absent as indicated by Whitworth (1958). Other diagnostic features are the larger length of the premolar row compared to the selenodont molar row and the presence of the dorcatherium fold in the lower molars. The genus Dorcatherium is also reported from Asia alongwith the Europe (Kaup, 1833; Lydekker, 1876; Matthew, 1929; Corbet and Hill, 1980, Farooq et al., 2008). Two species of genus Dorcatherium are known from Chinji formation i.e. D. majus and D. minus. D. minus. Isolation of protocone from hypocone, the presence of styles, the anterior median rib is stronger than the posterior one, the absence of entostylid are the characters of genus Dorcatherium while finely rugose enamel, comparatively weak mesostyle and well developed lingual cingulum is the specific character in the upper molars of D. minus (Colbert, 1935). In American museum collection two variations of D. minus can be observed: 1. Most of the upper molars have smaller mesostyle while the internal cingulum is strongly developed. 2. Few molars have heavy mesostyle while the cingulum is poorly developed (Colbert, 1935). The specimen under study (PUPC 08/90) is a left M1. It is not so large enough that may be included in the species D. majus. According to size and morphologically the specimen under study agrees with the D. minus. It compares favorably with AMNH 19517, AMNH 29856 and GSI B 195 and has similar antero-posterior length and crown width and W/L indices with already studied specimens by Farooq (2006) and Farooq et 81 al. (2007 a,b and 2008) (Table 6). The slight difference in the measurements is due to the individual variations. It is, therefore, concluded that on the basis of features like tooth morphology, over all contour of the tooth, size of the tooth, enamel constriction and development of the different crown structures, that specimen under study is referred to the genus Dorcatherium and the species D. cf. minus, however more material is needed for precise species identification. Bivariate scatter graph showing comparison of the studied specimens with the type specimens and previously studied specimens is also provided (Plot 12).

82

Family: GIRAFFIDAE Gray, 1821 Subfamily: GIRAFFINAE Zittel, 1893

Genus: Giraffa Brisson, 1762

Type Species: Giraffa camelopardalis Linnaeus 1758

Included Species: 1. G. camelopardalis Linnaeus 1758 2. G. sivalensis Falconer and Cautley 1843 3. G. punjabiensis pilgrim 1910 4. G. priscilla Mathew 1929

Distribution: The genus Giraffa is known from Upper, Middle and Lower Siwaliks (Pilgrim, 1910; Mathew, 1929 and Colbert, 1935). It is also known from Africa (Simpson, 1945).

Diagnosis: Medium sized giraffids with extremely elongated neck and limbs, skull with a moderately larger post orbital development; basicranial and basifacial axes inclined at a small angle. Paired parieto-frontal bony processes of small size and a median naso-frontal protuberance in both sexes; in some species paired occipital processes. A pre-lachrymal vacuity is present. Dentition very brachyodont, enamel very rugose, enamel folds pertaining deeply into the crown and enamel islands not formed until a late wear stage, lobes very oblique to the axis of the tooth. External ribs of upper teeth very strongly marked, outgrowths of enamel from the crescents into the central cavity. Length is not in excess of breadth, tubercles variable, but generally rudimentary, cingulum absent. Lower molars not elongated, tubercles in external valleys variable but a large one always present in M1 and M3, cingulum absent (Colbert, 1935).

83 Giraffa priscilla Mathew, 1929

Type specimen: GSI B 511, a left M3.

Locality: Chinji, Salt range, Chakwal district, Punjab, Pakistan (Colbert, 1935).

Stratigraphic range: This species is only found in the Lower Siwaliks (Chinji formation) of Pakistan.

Diagnosis: Distinguished from Giraffokeryx by the broad and more brachyodont teeth, prominent styles (especially metastyle). Prominent anterior rib; in M3 the more oblique- set inner crescent, broad third lobe with strong accessory basal cusps in front of it, as well as shorter crown (Colbert, 1935).

Specimens Examined: PUPC 08/29, an isolated right second lower molar and PUPC

08/10, left mandibular ramus having M1-2.

Locality: Dhok Bun Ameer Khatoon, Chakwal district, Punjab, Pakistan.

Stratigraphic range: Lower Siwaliks (Chinji Formation).

Description

PUPC 08/10 (Figure 21) The specimen under study consists of left mandibular fragment. Specimen is very well preserved. The mandible is missing anteriorly as well as posteriorly and having M1-2. The ascending ramus is missing in the specimen under description. It is moderately thick transversely and deep vertically. The depth below M1 is 34 mm and the width below M1 is 21 mm. Actual preserved mandible is 70 mm.

M1 It is in an early stage of wear. It is hypsodont and narrow crowned tooth (Table 7). Enamel is moderately thick and very rugose. The rugosity is more prominent on the 84 buccal side as compared to the lingual side. All the conids are well developed and well preserved except entoconid which is broken on the lingual side. Mesostylid is well developed. It is hypsodont tooth and has prominent styles (especially metastyle). Prominent anterior rib is present. The posterior central cavity is narrow and is formed between the hypoconid and entoconid. The transverse diameter of tooth is smaller as compared to the antero-posterior diameter. The inner cusps are slightly higher vertically as compared to the outer cusps. The protoconid is well developed and is present at the anterior buccal side. It is slightly higher but backward than the hypoconid and is connected with the metaconid through a thin enamel layer. Metaconid is present at the anterior lingual side of the tooth. It is pointed in the middle due to sloping ridges. On the lingual and posterior side of the tooth entoconid is present. Hypoconid is well developed and V shaped. The mesostylid is more developed as compared to metastylid. Metastylid is more developed as compared to other species. The median rib of metaconid and entoconid are moderately developed. The anterior central cavity is wider than posterior central cavity and both cavities are filled with shale and sand stone.

M2 It is well preserved and inserted into left mandible. It is in an early stage of wear. It is hypsodont tooth with well prominent styles (especially metastyle). Its anterior rib is prominent. It is narrow crowned tooth (table 7). Its enamel is moderately thick and very rugose. Rugosity is more prominent on the buccal side as compared to the lingual side. The median basal pillar is entirely absent. The anterior half of the tooth is wider as compared to the posterior half. The transverse diameter of the tooth is smaller as compared to the antero-posterior diameter. The major conids are strongly developed. The inner conids are slightly higher vertically as compared to the outer conids. The protoconid is well developed. It is located at the anterior buccal side of the tooth and is slightly higher than the hypoconid and is connected with metaconid through a thin enamel layer. The length of the anterior limb of protoconid is greater as compared to the posterior limb. The metaconid is well developed and is pointed in the middle with antero-posterior sloping ridges. Mesostylid is well developed while metastylid is moderately developed. The upper portion of metaconid is broken. The posterior limb of the metaconid touches the anterior limb of entoconid. The entoconid is located on lingual side and at the posterior side of the metaconid and is pointed in the middle with sloping ridges. The hypoconid is well preserved and crescent shaped. The stylids are more prominent at the 85 summit of crown and less distinct to the base of the tooth. Deep transverse valleys are present between cuspids.

PUPC 08/29 (Figure 20) It is an isolated right second lower molar and is well preserved. It is in an early stage of wear. Major conids are well developed and well preserved with stylids more prominent at the summit of crown and less distinct to the base of the tooth and transverse valleys filled with sand stone, broad in the middle and narrow antero-posteriorly.

86

A

B

C

Figure 20: Giraffa priscilla, PUPC 08/29, right second lower molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

87

A

B

C

Figure 21: Giraffa priscilla, PUPC 08/10, left mandibular ramus having M1-2, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

88

Table 7: Measurements (in mm) of molar teeth referred to Giraffa priscilla Mathew, 1929.

Number Position Length Width W/L

PUPC 08/10 M1 26 16 0.61

PUPC 08/10 M2 27.5 17 0.62

PUPC 08/29 M2 26 19 0.73

89

Lower M2 19.5

19

18.5 PUPC 08/10

18 PUPC 08/29 Length 17.5

17

16.5 25.5 26 26.5 27 27.5 28 Width a

Lower M1 and M2 19.5

19

18.5

18 PUPC 08/10 17.5 PUPC 08/10

Length PUPC 08/29 17

16.5

16

15.5 25.5 26 26.5 27 27.5 28 Width b Plot 13 a-b: Bivariate scatter graphs showing comparison of the studied specimens.

W/L index

0.8

0.7

M1 M2

0.6

0.5 1 2 3 4 5 6 7 8 9 10 11 12

Plot 14: Width/length indexes of cheek teeth of G. priscilla. 90

Discussion

In genus Giraffa styles are strong, median ribs are well pronounced and crown is broader while in Giraffokeryx styles are weak, median ribs are absent and crown is narrow. In lower molars of genus Giraffa stylids are present and median ribs stronger while in Giraffokeryx stylids are absent (Pilgrim, 1910) and median ribs are weaker. Giraffa punjabiensis is smaller than G. sivalensis and G. camelopardalis. Its upper premolars are relatively small and narrow while lower molars are long and lobes of molars set less obliquely to axis of jaw than in the recent giraffe. G. sivalensis is a larger species than modern giraffe. Its posterior half of last upper molar in the fossil form is reduced. Camelopardalis affinis is like G. sivalensis but larger in size to the modern giraffe. In G. priscilla broader and more brachyodont teeth, prominent styles (especially metastyle is prominent), prominent anterior rib. Parastyle is comparatively less developed as compared to the meso and metastyle that are well developed. Posterior median rib is also present. In the light of above mentioned characteristics it is evident that the specimens under study belong to genus Giraffa and all the above mentioned characters of the specimen under discussion are comparable with type material GSI B 492 (described by Matthew, 1929) of the species G. priscilla. Therefore it is concluded that, on the basis of above mentioned characters, the overall contour of the teeth, size of the teeth, enamel constriction and development of the different crown structure, specimens are referred to species G. priscilla. Graphs showing width/length indexes of cheek teeth of G. priscilla and comparison of the studied specimens are also provided (Plot 13 a-b and 14).

91

Genus Giraffokeryx Pilgrim, 1910

Type Species: Giraffokeryx punjabiensis Pilgrim, 1910

Included Species 1- G. punjabiensis Pilgrim, 1910 2- G. chinjiensis Sarwar, 1990

Distribution: The genus Giraffokeryx is only known from the Lower to Middle Siwaliks (Pilgrim, 1910 and Colbert, 1935).

Diagnosis: Giraffokeryx is a medium sized giraffid with four horns, two being at the anterior extremities of the fronto-parietal region. The posterior horn is overhanging the temporal fossa. Limbs and feet most probably have medium length. The teeth are brachydont with rugose enamel, as in other giraffids (Pilgrim, 1910).

Giraffokeryx punjabiensis Pilgrim, 1910

Type Specimen: Lectotype GSI 502, a third molar M3of right maxilla

Locality: Chinji and Vicinity, slat range Punjab.

Stratigraphic range: Lower Siwaliks, Chinji formation and lower portion of the Middle Siwaliks.

Diagnoses: G. punjabiensis is larger than G. chinjiensis. The upper molars are also comparatively larger and more hypsodont than those of G. chinjiensis. The para-, meso- and metastyle are not well pronounced. Median outer rib of paracone is present but weak. Teeth are brachyodont with rugose enamel. The median basal pillar is absent and the median ribs are moderately developed. The cingulum is slightly developed on the anterior side of the protocone. 92

Distribution: The species is known from the Kamlial and Chinji Formation (Lower Siwaliks) and the Nagri Formation of the Middle Siwaliks (Pilgrim, 1910; Colbert, 1935). The Lower Siwalik localities are Kamlial and Chinji, Attock and Chakwal districts respectively, within the Punjab Province of Pakistan. The locality of the Nagri zone is in the Nathot, Jhelum district of Punjab, Pakistan.

Specimens Examined: PUPC 08/35, an isolated left second upper premolar; PUPC 08/97, an isolated right second upper premolar; PUPC 08/33, an isolated right third upper premolar; PUPC 08/34, PUPC 08/43 and PUPC 08/98, isolated left third upper premolars; PUPC 08/26, an isolated left fourth upper premolar; PUPC 08/96, an isolated right fourth upper premolar; PUPC 08/31 and PUPC 08/105, isolated right first upper molars; PUPC 08/113, an isolated left first upper molar; PUPC 08/99, an isolated left second upper molar; PUPC 08/18 and PUPC 08/28, isolated right second upper molars; PUPC 08/117, an isolated right third upper molar; PUPC 08/38, an isolated right third lower premolar; PUPC 08/94 and PUPC 08/95, isolated left third lower premolars; PUPC 08/45, an isolated left first lower molar; PUPC 08/13, an isolated right lower second molar; and PUPC 08/11, an isolated right third lower molar. Locality: Dhok Bun Ameer Khatoon, Chakwal district, Punjab, Pakistan. Stratigraphic range: Lower Siwaliks (Chinji Formation).

Description Upper dentition P2 The specimens under study (PUPC 08/35 and PUPC 08/97) (Figures 22, 23) are isolated upper left and right second premolars respectively. They display excellent preservation, and are in the middle stage of wear. Both are low and narrow crowned. The anterio-posterior length is more than the transverse width of the teeth. The enamel is moderately dense, with a thickness of 1.5 mm, and is rugose, with the rugosity being more prominent on the lingual side, when compared to the buccal side, of the tooth. A cingulum is not developed. The central cavity of the tooth is well developed and narrow; furthermore, it is deep and located more toward the anterior of the tooth. It is slightly taller in the centre of the tooth than at its anterior and posterior ends. The parastyle is well developed whilst the mesostyle is moderately developed on the buccal side. The 93 metastyle is weakly developed. Between the parastyle and mesostyle, a faint layer of cement can be observed.

P3 PUPC 08/33 is an isolated right and PUPC 08/34, PUPC 08/43 and PUPC 08/98 (Figures 24-27) are isolated left third premolars. The teeth are in an early stage of wear. They are well preserved except PUPC 08/34, which is broken anteriorly; the protocone and metacone are absent. PUPC 08/43 and PUPC 08/98 display more wear than PUPC 08/33 and PUPC 08/34. Tooth crowns are narrow. Anterio-posterior length is more than the transverse width. The enamel is rugose and equally evident around the entirety of the tooth. All major cusps are well preserved. Proto- and paracones are wider while meta- and hypocones are shorter. A cingulum is moderately developed on the anterior side of the right P3 (PUPC 08/33), while it is faint and can only be observed on the lingual side of PUPC 08/34. There is no definite boundary between the major cones. The central cavity is formed by the union of major cusps, and is narrow and deep in the centre, while being shallow at the anterior and posterior ends. Para- and mesostyles are moderately developed and a groove is formed between them. P4 PUPC 08/26 and PUPC 08/96 (Figures 28, 29) are isolated left and right fourth upper premolars respectively. PUPC 08/26 is in an excellent state of preservation, whereas PUPC 08/96 is damaged. They are low and broad crowned. Anterio-posterior length is less than the transverse width of the teeth. Teeth are in the middle stages of wear. The enamel is rugose and moderately thick (2 mm). The rugosity is more prominent on the lingual side, compared to the buccal side in PUPC 08/26. In PUPC 08/96, the enamel is broken and dentine is visible on the posterior and lingual sides. A cingulum is not developed. The central cavity is shallow and extended more towards the anterior side and turns back towards the parastyle. A small amount of cement can be observed between the parastyle and mesostyle, which are both moderately developed, whilst the metastyle is well developed. The major cusps are moderately preserved. Crown height is greater at the buccal and lingual sides, compared to the smaller anterior and posterior sides. The outer cusps of the teeth are taller than the inner cusps. The anterio-posterior length of the buccal side is more than the lingual side of the tooth. The metacone and hypocone are slightly wider, whilst the protocone and paracone are narrow. Styles are weakly developed. 94 Parastyle, mesostyle and metastyle are not prominent. The median outer rib of the paracone is weak. Median ribs are present but less pronounced.

M1 PUPC 08/31 and PUPC 08/105 (Figures 30, 31) are isolated first molars of the right maxilla. PUPC 08/113 (Figure 32) is an isolated left first molar. Molars are in an excellent state of preservation and are in the middle stages of wear. The teeth being described are extremely brachyodont and narrow crowned. The enamel is moderately thick and is very rugose. The rugosity is more evident on the lingual side compared to the buccal side. Median basal pillar is absent. A weakly developed cingulum is present on the posterior side of the teeth. The protocone is broken in PUPC 08/31 and PUPC 08/105 while the other major cusps are well preserved; in PUPC 08/113 (Figure 32) all the cones are well preserved. The outer cones are slightly taller than the inner ones. The paracone is present on the anterior side of the teeth. The middle of the paracone is well developed and slightly broader, having two sloping anterio-posteriorly running ridges. The anterior ridge of the paracone is united with the parastyle while the posterior ridge touches the anterior ridge of the metacone. Parastyle of PUPC 08/113 is broken. Metacone is well developed. The height of the metacone is equal to the paracone and points to the middle of the tooth with two sloping ridges. The hypocone is crescentic in shape and has anterior and posterior limbs. The anterior limb is relatively small compared to that of the posterior. The parastyle and mesostyle are well developed while the metastyle is not. The anterior median rib is well developed while the posterior median rib is less so. Anterior and posterior central cavities are present. The anterior central cavity is narrow and not as shallow as the central cavity of the posterior.

M2 PUPC 08/18 and PUPC 08/28 (Figures 34, 35) are isolated second molars of the right maxilla, whilst PUPC 08/99 (Figure 33) is an isolated left second molar. The teeth of the right maxilla are in an excellent state of preservation; however, the buccal side of PUPC 08/99 is damaged. PUPC 08/18(Figure 34) is in an early stage of wear while no wear is observed in PUPC 08/28. Teeth are brachyodont and narrow crowned. The enamel is very thick and rugose. The wrinkles are more prominent on the lingual side compared to the buccal side. The median basal pillar is absent. The cingulum is well developed on the anterior side of the teeth. It is prominent on the anterior side of 95 protocone and is moderately developed around the hypocone. Comparatively, it is more developed on the anterior of the protocone compared with that of the posterior. The major cones are very well developed and the outer cones are slightly taller than the inner cones. A well developed protocone is present at the anterior lingual side of the teeth and is rounded in the middle. It has two limbs, one anterior and the other posterior, with the anterior limb being larger than the posterior. The paracone is pointed toward the middle of the tooth and is present at the anterio-buccal side of the teeth. From a posterior viewpoint, it is connected to the metacone, which is well developed. The metacone is slightly taller than the paracone. The mesostyle is well developed and the anterior median rib is more prominent than that of the posterior. The hypocone is present on the posterio- lingual side of both teeth and is well developed, having no connection between the hypocone and protocone. The parastyle is prominent. The paracone and metacone are joined to form an isolated pillar like structure known as the mesostyle. The parastyle, metastyle and mesostyle are absent in PUPC 08/99 because it’s buccal side is deteriorated.

M3 PUPC 08/117 (Figure 36) is an isolated third molar of the right maxilla. The tooth is in an excellent state of preservation, and is in the middle stages of wear. It is brachyodont and narrow crowned. The enamel is very thick and rugose. The wrinkles are more prominent on the lingual side compared to those of the buccal side. The median basal pillar is absent. The cingulum is well developed on the anterior side of the tooth. The major cones are very well developed and the outer cones are slightly taller than the inner cones. A well developed protocone is present on the anterio-lingual side of the tooth and is rounded in the middle. The anterior limb is larger than the posterior. The paracone is medially pointed and is present at the anterio-buccal side of the tooth. At the posterior side it is connected to the metacone which is well developed. The metacone is slightly taller than the paracone. The mesostyle is well developed and broken. The anterior median rib is more prominent than that of the posterior. The hypocone is present on the posterio-lingual side of the tooth and is well developed, having no connection with the protocone. The parastyle is prominent. The impression mark is absent on the posterior side of the tooth which confirms that the specimen under study is the last molar. 96 Lower dentition

P3 PUPC 08/38 (Figure 37) is an isolated right third lower premolar, and PUPC 08/94 and PUPC 08/95 (Figures 38-39) are isolated left third lower premolars. PUPC 08/95 is a well preserved tooth, whilst PUPC 08/38 and PUPC 08/94 are broken anteriorly. The teeth are in the middle stages of wear. Dentine can be seen on all conids. They are hypsodont and narrow crowned. The enamel is thick and rugose; the rugosity is more prominent on the outer side. The paraconid is fairly distinct and separated from the parastylid by a wide furrow. The paraconid and the metaconid are not fused and a deep valley is present between them. The metaconid is higher than the paraconid which is growing backward, enclosing an elongated and transverse valley together with the entoconid. The entostylid is weakly developed. The hypoconid is projected latterly with a deep valley in front of it.

M1 PUPC 08/45 (Figure 40) is a lower left isolated first molar and is in the early stages of wear. It is hypsodont and narrow crowned. The enamel is moderately thick and very rugose. The rugosity is more prominent on the buccal side, compared to that of the lingual side. All conids are well developed. The entoconid is slightly higher than the hypoconid. The mesostylid is well developed. The posterior central cavity is narrow and is formed between the hypoconid and entoconid. The transverse diameter of the tooth is smaller than the anterio-posterior diameter. The inner cusps are slightly taller than the outer cusps. The protoconid is well developed and present on the anterio-buccal side. It is slightly higher than the hypoconid and is connected with the metaconid through a thin enamel layer. The metaconid is present at the anterio-lingual side of the tooth. The entoconid is present on the lingual and posterior side of the tooth. The entoconid is pointed in the middle due to sloping ridges. The hypoconid is well developed and V- shaped. The mesostylid is more developed than the metastylid. The median ribs of the metaconid and entoconid are moderately developed. The anterior central cavity is wider than the central cavity of the posterior.

M2 PUPC 08/13 (Figure 41) is a lower right isolated second molar and is in the early stages of wear. It is hypsodont and narrow crowned. The enamel is moderately thick and 97 very rugose. The rugosity is more prominent on the buccal side compared to that of the lingual side. All conids are well developed. The entoconid is slightly taller than the hypoconid. The mesostylid is well developed. The posterior central cavity is narrow and is formed between the hypoconid and entoconid. The transverse diameter of the tooth is smaller compared to the anterio-posterior diameter. The inner cusps are slightly taller compared to the outer cusps. The protoconid is well developed and is present at the anterio-buccal side. It is slightly higher than the hypoconid and is connected with the metaconid through a thin enamel layer. The metaconid is present on the anterio-lingual side of the tooth. The entoconid is present on the lingual and posterior side of the tooth and is pointed in the middle due to sloping ridges. The hypoconid is well developed and V shaped. The mesostylid is more developed compared to the metastylid. The median ribs of the metaconid and entoconid are moderately developed. The anterior central cavity is wider than the central cavity of the posterior.

M3 PUPC 08/11 (Figure 42) is an isolated third molar of the right mandible and the tooth is embedded in the jawbone. It is well preserved and is in the early stage of wear. It is extremely narrow crowned. Its enamel is moderately thick and very rugose. The median basal pillar is entirely absent. The cingulum is weakly developed on the anterior side of the protoconid. The anterior half of the tooth is wider than the posterior half. The transverse diameter of the tooth is smaller compared to the anterio-posterior diameter. The major conids are strongly developed. The inner conids are slightly taller than the outer ones. The protoconid is well developed. It is located at the anterio-buccal side of the tooth and is slightly higher than the hypoconid and is connected with the metaconid through a thin layer of enamel. The length of the anterior limb of the protoconid is greater compared to the posterior limb. The metaconid is well developed and is pointed in the middle with anterio-posterior sloping ridges. The mesostylid is well developed, whilst the metastylid is weaker. The posterior limb of the metaconid touches the anterior limb of the entoconid. The entoconid is located on the lingual side and at the posterior side of the metaconid and is pointed in the middle with sloping ridges. A strong and well developed talonid is present on the posterior side of the tooth. The posterior end of the entoconid is joined with the talonid. The hypoconid is well preserved and crescent shaped. The stylids are more prominent at the summit of the crown and less distinct at the base of the tooth. Deep transverse valleys are present between the cuspids. 98

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Figure 22: G. punjabiensis, PUPC 08/35, isolated left second upper premolar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

99

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Figure 23: G. punjabiensis, PUPC 08/97, isolated right second upper premolar. A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

100

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Figure 24: G. punjabiensis, PUPC 08/33, isolated right third upper premolar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

101

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Figure 25: G. punjabiensis, PUPC 08/34, isolated left third upper premolar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

102

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Figure 26: G. punjabiensis, PUPC 08/43, isolated left third upper premolar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

103

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Figure 27: G. punjabiensis, PUPC 08/98, isolated left third upper premolar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

104

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Figure 28: G. punjabiensis, PUPC 08/26, isolated left fourth upper premolar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 105

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Figure 29: G. punjabiensis, PUPC 08/96, isolated right fourth upper premolar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 106

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Figure 30: G. punjabiensis, PUPC 08/31, isolated right first upper molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

107

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Figure 31: G. punjabiensis, PUPC 08/105, isolated right first upper molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

108

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Figure 32: G. punjabiensis, PUPC 08/113, isolated left first upper molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 109

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Figure 33: G. punjabiensis, PUPC 08/99, isolated left second upper molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

110

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Figure 34: G. punjabiensis, PUPC 08/18, isolated right second upper molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 111

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C Figure 35: G. punjabiensis, PUPC 08/28, isolated right second upper molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

112

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Figure 36: G. punjabiensis, PUPC 08/117, isolated right third upper molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

113

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Figure 37: G. punjabiensis, PUPC 08/38, isolated right third lower premolar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 114

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Figure 38: G. punjabiensis, PUPC 08/94, isolated left third lower premolar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 115

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Figure 39: G. punjabiensis, PUPC 08/95, isolated left third lower premolar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 116

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Figure 40: G. punjabiensis, PUPC 08/45, isolated left first lower molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 117

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Figure 41: G. punjabiensis, PUPC 08/13, isolated right second lower molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

118

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Figure 42: G. punjabiensis, PUPC 08/11, isolated right third lower molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 119 Table 8: Comparative measurements (mm) of the cheek teeth referred to G. punjabiensis Pilgrim, 1910 (AMNH taken from Colbert, 1935).

Number Position Length Width W/L

AMNH 19475 P2 22 19 0.86 PUPC 08/35 P2 22 19 0.86 PUPC 08/97 P2 20.5 18 0.88 AMNH 19475 P3 20.5 20 0.98 PUPC 08/33 P3 23 21.5 0.93 PUPC 08/34 P3 23 21 0.91 PUPC 08/43 P3 16 13 0.81 PUPC 08/98 P3 23 20 0.87 AMNH 19475 P4 7.5 21 1.2 AMNH 19330 P4 17 23 1.4 PUPC 08/26 P4 18.5 21 1.1 PUPC 08/96 P4 13.5 19 1.4 AMNH 19334 M1 24 24 1 AMNH 19311 M1 25.5 25 0.98 PUPC 08/31 M1 25.5 25 0.98 PUPC 08/105 M1 24 23 0.96 PUPC 08/113 M1 26 26 1 AMNH 19472 M2 27 25 0.93 AMNH 19611 M2 26 27 0.96 AMNH 19320 M2 29 28.5 0.98 PUPC 08/18 M2 28.5 27 0.95 PUPC 08/28 M2 30 29 0.97 PUPC 08/99 M2 26 26 1 AMNH 19325 M3 27.5 28 1.1 AMNH 19327 M3 23 23.5 1.2 AMNH 19472 M3 25 25 1 AMNH 19475 M3 24.5 26 1.1 PUPC 08/117 M3 32.5 33 1.1 AMNH 19587 P3 20.5 12 0.86 120

Number Position Length Width W/L AMNH 19849 P 22 11 3 0.5 PUPC 08/38 P 20 12 3 0.6 PUPC 08/94 P 21 10.5 3 0.5 PUPC 08/95 P 22 13 3 0.59 AMNH 19849 M 20 14.5 1 0.73 AMNH 19587 M 24 14 1 0.58 AMNH 19593 M 24 16 1 0.67 PUPC 08/45 M 19 12 1 0.63 AMNH 19849 M 22 16 2 0.72 AMNH 19587 M 25 17 2 0.68 AMNH 19324 M 27 19 2 0.71 PUPC 08/13 M 28 18 2 0.64 AMNH 19449 M 35 15.5 3 0.44 AMNH 19587 M 37 17 3 0.46 AMNH 19324 M 38 17 3 0.45 PUPC 08/11 M 39 16 3 0.41 121

Upper P2 22.5 22 21.5 AM NH 19475 PUPC 08/35

21 PUPC 08/97 Length 20.5 20 17.5 18 18.5 19 19.5 Width

a Upper P3 25 AM NH 19475 20 PUPC 08/33 PUPC 08/34 15 PUPC 08/43 PUPC 08/98

Length 10 5 0 10 15 20 25 Width

b Upper P4 20

15 AM NH 19475 AM NH 19330 10 PUPC 08/26

PUPC 08/96 Length 5

0 14 18 22 26 Width

c 122

Upper M1 26.5 26 AM NH 19334 25.5 AM NH 19311 25 PUPC 08/31 PUPC 08/105 Length 24.5 PUPC 08/113 24 23.5 22 23 24 25 26 27 Width

d Upper M2 31

30 AM NH 19472 AM NH 19611 29 AM NH 19320 28 PUPC 08/18 PUPC 08/28 Length 27 PUPC 08/99 26 25 24 26 28 30 Width

e Upper M3

35 30 25 AM NH 19325 AM NH 19327 20 AM NH 19472 15 AM NH 19475 Length PUPC 08/117 10 5 0 20 25 Width 30 35

f 123

Lower P3 22.5 AM NH 19587 22 AM NH 19849 PUPC 08/38 21.5 PUPC 08/94 21 PUPC 08/95

Length 20.5 20 19.5 8 10 12 14 16 Width

g Lower M1 30 AM NH 19849 25 AM NH 19587 20 AM NH 19593 15 PUPC 08/45

Length 10 5 0 10 12 14 16 18 Width

h Low er M2 30 AM NH 19849 20 AM NH 19587 AM NH 19324 PUPC 08/13

Length 10

0 15 16 17 18 19 20 Width

i 124

Lower M3 40 39 AM NH 19449 38 AM NH 19587 37 AM NH 19324 PUPC 08/11

Length 36 35 34 15 15.5 16 16.5 17 17.5 Width

j

Plot 15a-j: Bivariate scatter graphs showing comparison of the studied specimens with the type specimens.

Upper dentition

35 AMNH 19475 (P2) PUPC 08/35 (P2)

PUPC 08/97 (P2) AMNH 19475 (P3) 30 PUPC 08/33 (P3) PUPC 08/34 (P3)

PUPC 08/43 (P3) PUPC 08/98 (P3) 25 AMNH 19475 (P4) AMNH 19330 (P4)

PUPC 08/26 (P4) PUPC 08/96 (P4) 20 AMNH 19334 (M1) AMNH 19311 (M1)

PUPC 08/31 (M1) PUPC 08/105 (M1)

Length 15 PUPC 08/113 (M1) AMNH 19472 (M2)

AMNH 19611 (M2) AMNH 19320 (M2) 10 PUPC 08/18 (M2) PUPC 08/28 (M2)

PUPC 08/99 (M2) AMNH 19325 (M3) 5 AMNH 19327 (M3) AMNH 19472 (M3)

AMNH 19475 (M3) PUPC 08/117 (M3) 0 0 5 10 15 20 25 30 35 Width

Plot 16: Plot shows the size variation in the upper dentition of G. punjabiensis

125

Lower dentition

20

18 AMNH 19587 (P3) 16 AMNH 19849 (P3) PUPC 08/38 (P3) PUPC 08/94 (P3) 14 PUPC 08/95 (P3) AMNH 19849 (M1) 12 AMNH 19587 (M1) AMNH 19593 (M1) 10 PUPC 08/45 (M1)

AMNH 19849 (M2) Length 8 AMNH 19587 (M2) AMNH 19324 (M2) 6 PUPC 08/13 (M2) AMNH 19449 (M3) 4 AMNH 19587 (M3) AMNH 19324 (M3) 2 PUPC 08/11 (M3)

0 0 10 20 30 40 50 Width

Plot 17: Plot shows the size variation in the lower dentition of G. punjabiensis

W/L index of upper dentition

1.6

1.4

1.2 P2 1 P3 P4 0.8 M1

0.6 M2 M3 0.4

0.2

0 1 2 3 4 5 6

Plot 18: Width/length indexes of the upper cheek teeth of G. punjabiensis

126

W/L index of Lower dentition

1

0.9

0.8

0.7

0.6 P3 M1 0.5 M2 0.4 M3

0.3

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0.1

0 1 2 3 4 5

Plot 19: Width/length indexes of the lower cheek teeth of G. punjabiensis 127 DISCUSSION

The specimens described are of typical giraffid type, which show basic features, such as the depth of enamel folds and the rugose sculpture of the enamel, and they are here referred to G. punjabiensis. Specimen PUPC 08/35 is a second upper left premolar and PUPC 08/97 is an isolated right P2. When they are compared with AMNH 19475 (identified as G. punjabiensis by Colbert, 1935), it is evident that the anterio-posterior length and crown width are identical in the case of PUPC 08/35 and compare favorably with PUPC 08/97. They are narrow crowned while AMNH 19475 is also narrow crowned. All the major cusps are well preserved with a grooved lingual side in the middle while the buccal side is spindle shaped. The anterio-posterior length of the teeth is greater than the transverse width (Table 8). Median ribs are missing while the styles are weakly preserved which indicate that specimens under study belong to G. punjabiensis. The specimens PUPC 08/33, PUPC 08/34, PUPC 08/43 and PUPC 08/98 are also smaller in size. On the basis of tooth size, contour and enamel constriction and structure of crown, they resemble, based on information given by Colbert, 1935, the species G. punjabiensis. The P3 measurements described here strongly resemble those given for AMNH 19475. They are all narrow crowned and their W/L ratio also resembles the type specimen. In G. punjabiensis the external folds are comparatively more developed in the premolars, and in the specimens under study the styles and median ribs are less pronounced which is a feature of G. punjabiensis. The specimens PUPC 08/26, PUPC 08/96 are fourth upper left and right premolars respectively, and when compared with AMNH 19475 and AMNH 19330, they resemble the known material of G. punjabiensis. All of the teeth are broad crowned and show the typical morphology of this species. Additionally, crown width is greater than the anterio-posterior length. Moreover, the anterio-posterior length, transverse width and W/L ratio also resemble the P4 of AMNH 19475 and AMNH 19330. In specimens PUPC 08/31, PUPC 08/105 and PUPC 08/113 the median ribs and styles are less pronounced and the measurements and tooth morphology resemble AMNH 19334 and AMNH 19311. The specimens presently under study, together with the M1 in the AMNH collections, are narrow crowned. PUPC 08/18 and PUPC 08/28 are right upper second molars while PUPC 08/99 is a left molar and is comparable to AMNH 19320, AMNH 19472 and AMNH 19611 128 (Table 8). PUPC 08/117 is a right upper third molar and is comparable to AMNH 19325, AMNH 19327, AMNH 19472 and AMNH 19475. They have similar anterio- posterior lengths, crown widths and are narrow crowned. In the specimens under study, the parastyle is more developed compared to the mesostyle and metastyle, while the posterior median rib is also missing (Colbert, 1935). PUPC 08/38, PUPC 08/94 and PUPC 08/95 are lower third premolars. Measurements exhibit a close relationship with the type specimens AMNH 19587 and AMNH 19849. These lower third premolars resemble, in terms of the structure of the cusps, cingula, rugosity of enamel, anterio-posterior length and development of styles, the type specimen. Their W/L index lies close to specimen AMNH 19849. The teeth are narrow crowned. The specimens under study show all the morphological features of the cheek teeth cited by Colbert (1935). In Giraffokeryx the styles are weak, median ribs are absent and the crown is narrow. Stylids are absent in the lower molars. The lower first molar, on the basis of cusp structure, rugosity of enamel, anterio-posterior length, and W/L index, compares favorably with the type specimen and other specimens present in the AMNH, i.e. AMNH 19849, AMNH 19587 and AMNH 19593. Specimen PUPC 08/13 is a lower right second molar. When it is compared with AMNH 19849, AMNH 19587 and AMNH 19324, it is evident that all teeth are narrow crowned and their anterio-posterior length and crown width resemble each other. It is also observed that the anterio-posterior length is greater than the crown width. Stylids are weakly developed in the specimen under study, which is a major character of the genus Giraffokeryx. The specimen PUPC 08/11 is a right lower third molar embedded in a piece of jaw bone, and it resembles AMNH 19324, AMNH 19587 and AMNH 19849 (identified as G. punjabiensis by Colbert, 1935) in terms of anterio-posterior length and crown width (Table 8). The tooth under study also resembles the typical morphology of G. punjabiensis. Slight differences in the measurements obtained are due to intraspecific variation. The stylids and median ribs are less pronounced in the specimen under study which is a character of G. punjabiensis. Therefore, all the specimens under study are being referred to G. punjabiensis (Table 8). Plots (15 a-j, 16, 17, 18 and 19) show comparison of the studied specimens with the type specimens, the size variation in the upper and lower dentition and width/length indexes of the upper and lower cheek teeth of G. punjabiensis. 129 Family BOVIDAE Gray, 1821 Subfamily ANTILOPINAE Gray, 1821 Tribe ANTILOPINI Gray, 1821

Genus: Gazella Blainville, 1816

Type species: Gazella dorcas Linneus, 1758.

Included species 1. G. dorcas Linneus, 1758 2. G. gazella Pallas, 1758 3. G. dama Pallas, 1766 4. G. subgutturosa Guldenstacdt, 1780 5. G. rufifrons Gray, 1846 6. G. deperdita Gervais, 1847

7. G. capricornis Wagner, 1848 8. G. atlantica Bourguignat, 1870 9. G. leptoceros Cuvier, 1882 10. G. setifensis Pomel, 1895 11. G. thomasi Pomel, 1895 12. G. gaudryi Schlosser, 1904 13. G. longicornis Andree, 1926 14. G. mytilinii Pilgrim, 1926 15. G. rodleri Pilgrim and Hopwood, 1928 16. G. gaudryi Bohlin, 1935 17. G. pilgrimi Bohlin, 1935a

18. G. dorcacoides Bohlin, 1935a

19. G. lydekkeri Pilgrim, 1937

20. G. sinensis Teilhard and Piveteau, 1938

21. G. blacki Teilhard and Trassaert, 1938

22. G. arista Bate, 1940

23. G. decora Bate, 1940

24. G. praethomsoni Arambourg, 1947 130 25. G. wellsi Cooke, 1949

26. G. janenschi Dietrich, 1950

27. G. hennigi Dietrich, 1950

28. G. stehlini Thenius, 1951

29. G. gracilior Cooke and Wells, 1956

30. G. vanhoepeni Cooke and Wells, 1956

31. G. tingitana Arambourg, 1957

32. G. pregaudryi Arambourg, 1959

33. G. padriensis Akhtar, 1992

Distribution: The genus Gazella is known from the Miocene and Pliocene of Eurasia and several Pleistocene localities in Africa (Gentry, 1966). It is abundantly found at the same time in Asia, Siwaliks and southern parts of Europe (Pilgrim, 1937, 1939; Akhtar, 1992; Khan, 2009a).

Diagnosis: Upper molars are hypsodont having prominent narrow styles, without basal pillars, central cavities are crescentic in shape, median ribs moderately strong in primitive forms while weak or absent in progressive forms, P2 longer as compared to P3 and P4. Lower molars with goat folds, small ectostylid, central cavities having fairly simple outline, ribs and stylids are moderately developed.

Gazella sp. Type Specimen: A skull and conjoined mandible (AMNH 19663).

Locality: Dhok Pathan, Chakwal district, Punjab, Pakistan.

Stratigraphic range: Lower and Middle Siwaliks.

Diagnosis: Upper molars hypsodont, without entostyle and basal pillars while enamel is moderately thick and rugose, styles narrow and strong, anterior median rib stronger as compared to the posterior one, have narrow and deep central cavities, premolar series slightly long. Lower molars are extremely hypsodont with small basal pillars, well 131 developed goat folds, central cavities fairly simple in their outline and moderately developed stylids and ribs.

Distribution: The species is known from the Dhok Pathan stage of the Middle Siwaliks (Pilgrim, 1937). It survived to the Late Pliocene and its most recent record is from the upper levels of the Dhok Pathan stage of the Middle Siwaliks (Akhtar, 1992) Chinji type locality, Lower Siwaliks (Khan, 2009a) and Dhok Bun Ameer Khatoon, Chakwal district, Punjab, Pakistan.

Specimens examined: PUPC 08/16, right mandibular ramus with M1-2 and PUPC 08/111, an isolated left second upper molar and PUPC 08/116, left mandibular ramus with M1-2.

Locality: Dhok Bun Ameer Khatoon, Chakwal district, Punjab, Pakistan.

Stratigraphic range: Lower Siwaliks (Chinji Formation).

Description

Lower dentition PUPC 08/16 (Fig. 43) and PUPC 08/116 (Fig. 44) are right and left mandibular ramii, and well preserved. Total length of M1-2 in left mandible is 51 mm while right one is 50 mm. The height of mandibular ramus below left M1 is 19 mm and right M1 is 18.5 mm and the width below left M1 is 12 mm and right M1 is 11 mm. The lower teeth are very well preserved, narrow crowned and in an early wear. A well developed goat fold is present anteriorly and it is somewhat heavy towards labially. A well developed median basal pillar is present in the transverse valley. The lingual conids are higher than those of labial ones. The protoconid is rounded in general appearance. The praeprotocristid is larger in antero-posterior length than the postprotocristid and continuous with the goat fold. In general appearance, the hypoconid looks to be V-shaped and less crescentic than the protoconid. It is forwardly directed. The metaconid is well developed and posterior to it, a pointed entoconid is present. The entoconid is high in the middle with praeentocristid and postentocristid sloping ridges. The metastylids and entostylids are well developed. The anterior median rib is more prominent than the posterior one. 132

Upper dentition M2: PUPC 08/111(Fig. 45) is well preserved and in an early wear. It is hypsodont and narrow crowned tooth (Table 9). The enamel is moderately thick and rugose. The rugosity is more prominent on the lingual side as compared to the buccal side. The central cavities are wide and deep. The praeprotocrista is continuous with the paracone. The parastyle is well developed. The mesostyle is very strong, well developed and directed toward anterior side. The metastyle is damaged at the tip. The hypocone looks like crescentic in shape and is not uniform in thickness. The praehypocrista is comparatively thin than that of the posthypocrista. The paracone is almost equal to the metacone in antero-posterior length. The anterior rib is more prominent as compared to the posterior one which is only visible towards the crown top. Median basal pillar is absent. 133

A

B

C

Figure 43: Gazella sp., UPC 08/16, a right mandibular ramus having M1-2. A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

134

A

B

C

Figure 44: Gazella sp., UPC 08/116, a left mandibular ramus having M1-2. A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 135

A

B

C

Figure 45: Gazella sp., UPC 08/111, a left second upper molar. A - buccal view, B- occlusal view, C - lingual view. Scale bar 10 mm.

136 Table 9: Comparative measurements of the cheek teeth of Gazella sp. in mm (millimeters). * The studied specimens. (AMNH taken from Colbert,1935).

Number Position Length Width W/L

PUPC 08/16* M1 13 8.5 65.4

M2 15.5 9 0.58

PUPC 08/116* M1 15 8.5 56.6

M2 18 9 0.5

AMNH 19663 M1 12.0 10 0.83

PUPC 84/133 M1 12.0 6.0 0.5

PUPC 84/67 M1 14.5 9.0 0.62

PUPC 04/2 M1 11.0 8.2 0.74

M2 14.0 8.7 0.62

AMNH 19663 M2 13.0 7.5 0.57

PUPC 83/684 M2 12.0 8.0 0.66

PUPC 84/133 M2 15.5 10.0 0.64

PUPC 02/37 M2 14.5 9.0 0.62

PUPC 08/111* M2 13 12.5 0.96

AMNH 19663 M2 13.5 11.5 0.85

PUPC 84/65 M2 18.0 17.3 0.96

PUPC 85/82 M2 14.0 10.0 0.71

PUPC 97/21 M2 12.0 12.0 1.0

PUPC 97/22 M2 13.5 13.0 0.96

PUPC 00/101 M2 13.0 11.0 0.84

137

Lower M1 12

10

PUPC 08/16* 8 PUPC 08/116* AMNH 19663

6 PUPC 84/133 Length PUPC 84/67 4 PUPC 04/2

2

0 0 5 10 15 20 Width a

Lower M2 12

10 PUPC 08/16*

8 PUPC 08/116* PUPC 04/2

6 AMNH 19663

Length PUPC 83/684

4 PUPC 84/133 PUPC 02/37

2

0 0 5 10 15 20 Width b

Upper M2 20

18

16 PUPC 08/111* 14 AMNH 19663 12 PUPC 84/65 10 PUPC 85/82

Length PUPC 97/21 8 PUPC 97/22 6 PUPC 00/101 4

2

0 0 5 10 15 20 Width c Plot 20a-c: Bivariates showing comparison of the studied specimens with the type specimens.

138

W/L Ratio

1.2

1

0.8

Upper M2 0.6 Lower M2 Lower M1

0.4

0.2

0 1 2 3 4 5 6 7 8

Plot 21: Width/length indexes of the cheek teeth of Gazella sp.

Lower M1, M2 and Upper M2 20 PUPC 08/16* PUPC 08/116* 18 AMNH 19663 PUPC 84/133 16 PUPC 84/67 PUPC 04/2 14 PUPC 08/16* PUPC 08/116* 12 PUPC 04/2 AMNH 19663 10 PUPC 83/684

Length PUPC 84/133 8 PUPC 02/37 PUPC 08/111* 6 AMNH 19663 PUPC 84/65 4 PUPC 85/82 PUPC 97/21 2 PUPC 97/22 PUPC 00/101 0 0 5 10 15 20 Width

Plot 22: Plot shows the size variation in the dentition of Gazella sp.

139 Discussion The specimens under study are square and tetratuberculate; these can be referred to some herbivorous mammals. Since the cusps are crescentic in outline, so it can be included in sub-order ruminantia of the order artiodactyla. The molars being smaller in size and enamel layer is finely rugose, can be referred to family Bovidae. The teeth are not so large that they can be included in the large Siwalik bovids. The general contour of the studied specimens, upper molar hypsodonty, without entostyle and basal pillars while enamel is moderately thick and rugose, styles narrow and strong, anterior median rib stronger as compared to the posterior one, have narrow and deep central cavities, premolar series slightly long. Lower molars are extremely hypsodont with small basal pillars, well developed goat folds, central cavities fairly simple in their outline, moderately developed stylids and ribs evidently prove the specimens inclusion in subfamily Antilopinae. This subfamily is characterized by many genera including the genus Gazella. A very distinct and common feature of the specimens under study and the type is the presence of a single deep furrow on the posterior side, generic features of the genus Gazella cited by Pilgrim (1939). The dimensions (Table 9) and the morphology of the studied material reveal all the features of the species Gazella sp. cited by Pilgrim in 1937. About the upper molars Pilgrim (1937) stated that the antero-posterior length is decidedly greater than the transverse diameter in the case of the last two molars, but MI is slightly quadrate. All the teeth under study show typical features of this species as stated by Pilgrim (1937), "on the outer side of the molars the anterior and median folds are of about equal strength and rather prominent while the posterior fold is weak. The median rib of the anterior lobe is strongly developed, where as the posterior lobe is weaker, but both are visible to the base of the crown. Median basal pillars are not visible". No basal pillar could be detected in the upper studied tooth. In case of lower molars Pilgrim (1937) stated that At the anterior end, both on the inner as well as on the outer side, each molar has a strong fold, called goat fold. It is more prominent on the outer than on the inner side. At the posterior end there is only a weak fold on the inner side and none at all on the outer side. PUPC 08/16, PUPC 08/111 and PUPC 08/116 present the same morphological features of the type specimen AMNH 19663 (Pilgrim, 1937) and specimens studied by Khan, 2008. The type specimen and the specimens under study all are narrow crowned. 140 In case of PUPC 08/111 antero-posterior length of specimen is smaller and transverse width is more than the type specimen while all other teeth are, however, slightly larger than the type specimen but this, in all probability, is within the range of individual variation. Plots (20a-c, 21 and 22) are bivariates showing comparison of the studied specimens with the type specimens, width/length indexes and the size variation in the dentition of Gazella sp. The studied material is referred to genus Gazella but more material is required to identify it up to species level and due to scarcity of material, Gazella sp. is attributed for the material under study.

141 Genus: Eotragus Pilgrim, 1939

Type Species: Eotragus sansaniensis (Lartet, 1851).

Generic Diagnosis: Small sized bovid with the buccal walls of the upper molars inclined, and obliquely situated teeth so that the buccal walls do not line up. The triangular shaped horn core, when examined laterally (in profile) is not symmetrical longitudinally, but rather has more bone mass at the base of the anterior region than the base of its posterior region. This results in a characteristically wide horn core base. In lateral view, the posterior edge is usually convex while the anterior one is concave. These characteristics make the horn core axis curve to project slightly forward. The cross section of the Eotragus horn core is slightly oval, often resulting in a weak anterior keel. The horn core is situated above the orbits on a medium-sized pedicle. The orbital rims appear to protrude in relation to the position of the horn core pedicles. The horn cores are inclined posteriorly about 40 in lateral view (Solounias et al., 1995).

Included Species 1- E. sansaniensis Lartet, 1851 2- E. cristatus Biedermann, 1873 3- E. haplodon Thenius, 1952 4- E. artenensis Ginsburg and Heintz, 1968 5- E. halamagaiensis Ye, 1989 6- E. noyei Solounias et al., 1995

Distribution: Spain, Central Europe, Libya, Kenya, Israel, China and Pakistan.

Stratigraphic range: Lower and Middle Siwaliks.

142

Eotragus sp.

Specimen Examined: PUPC 08/100, a right mandibular ramus having P3-M3. PUPC

08/101, left mandibular ramus having M1-3 and PUPC 08/114, an isolated right first lower molar.

Locality: Dhok Bun Ameer Khatoon, Chakwal district, Punjab, Pakistan.

Stratigraphic range: Lower Siwaliks (Chinji Formation).

Description PUPC 08/100 (Figure 46) is right mandibular ramus having well preserved

P3- M3. The length of the preserved mandibular ramus is 71mm while the depth below the M2 is 16 mm and the M3 is 17 mm. The thickness of ramus below M2 is

9 mm and M3 is 10 mm. The teeth present a middle stage of wear. Specimen PUPC

08/101 (Figure 47) is a left mandible fragment having M 1-3. It is in an early stage of wear and seems to belong to a young individual. PUPC 08/114 (Figure 48) is an isolated right first lower molar. The tooth is very well preserved and is in an early stage of wear.

The P3 is well preserved with thick and shinny enamel. The tooth is narrow crowned and a layer of cement is prominent at the base of the crown. The median basal pillar is absent. All the conids of the tooth are well preserved.

P4 is narrow crowned with a conical structure and part of the right mandible. It is damaged on the lingual side while all other portions are well preserved. Its buccal side is also more rugose as compared to the lingual side which has quite thick and shinny enamel. Layer of cement is also prominent at the base of the crown. In the molar series of PUPC 08/100 and PUPC 08/101, the principal conids are well preserved and prominent. The lingual conids are very similar to each other in their general appearance and same is the case with the buccal ones. The lingal conids are broad, vertically higher and the buccal conids are roughly V- shaped. The lingual conids are broad in the middle while narrow antero- posteriorly. All the crown features are preserved and the teeth show the confluence of the metacristid, the protocristid and the postmetacristid. The median ribs and the mesostylid are weakly developed, whereas the metastylid are well developed. The anterior median rib is more prominent and strong than that of posterior one. A 143 small goat fold connected with the prastylid forms a transverse flange. A median basal pillar is present in the transverse valleys on the buccal sides of molars. The enamel is rugose and this rugosity is more prominent on the buccal side of the teeth while the cingulum is absent.

Well preserved M1-3 teeth are the part of both right and left mandibular ramii. The hypoconid is forwardly directed as compared to the protoconid. A thick layer of cement is present at the base of the right M 1-3. Central cavities are well preserved and these are narrow in the middle while broad antero-posteriorly. The first molar in PUPC 101 can be firmly determine as an M1, because of the typical rounded shape of the anterior contact facet for the premolar and the wider anterior part of the tooth comparatively to the posterior part of the tooth (Rossner, 1995) and trace of anterior cingulum is present in it. The second molars are larger than the first molars with anterolingual and anterolabial folds present, the basal pillar is weaker, and metastylid is conspicuous. In right third molar median basal pillar is hardly visible because of the thick layer of cement. It is broad at the base while narrow towards the crown. The central cavities are well preserved and these are narrow in the middle while broad antero-posteriorly. The crown of M2 is higher than that of M3. In M3, a well- developed talonid is present and its height is comparatively lower than the other major conids. It is opened anteriorly and roughly circular posteriorly. 144

A

B

C

Figure 46: Eotragus sp., PUPC 08/100, right mandibular ramus having P3-M3, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm.

145

A

B

C

Figure 47: Eotragus sp., UPC 08/101, left mandibular fragment with M1-3, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 146

A

B

C

Figure 48: Eotragus sp.,PUPC 08/114, a right first lower molar, A - buccal view, B - occlusal view, C - lingual view. Scale bar 10 mm. 147 Table 10: Comparative measurements (in mm) of the cheek teeth of Eotragus sp. (data taken from Alfarez et al., 1980; Solounias et al., 1995; Khan et al., 2008a, 2009). *The studied specimens. ** Eotragus sp. (large size) Khan et al., 2008a.

Number Position Length Width W/L

PUPC 08/100* P3 11.5 6 0.52 P4 11.5 7 0.61 M1 11 9 0.82 M2 15 10 0.67 M3 21 10 0.48 PUPC 08/101* M1 13 10 0.77 M2 15 10.5 0.7 M3 20 10 0.5 PUPC 08/114* M1 13.5 8.5 0.63 PUPC 04/24 M1 8.0 5.0 0.62 M2 8.0 5.6 0.7 PUPC 05/11 M2 8.7 4.6 0.52 M3 11.7 4.3 0.36 CO-489 M1 9.4 6.2 0.65 CO-490 M2 11.6 7.9 0.69 CO-491 M3 14.2 7.0 0.49 PC-GCUF 08/01 M3 11.0 4.84 0.43 PUPC 69/272 ** M1 13.5 8.0 0.59 M2 13.5 8.0 0.59

Table 11: Dental measurements of different species of the genus Eotragus (in mm) taken from Ye, 1989.

Species Position Length Width W/L

E. halamagaiensis M1 10.3 7.3 0.71

M2 13.3 8.2 0.62

M3 17.3 7.3 0.42

E. artenensis M2 11.5 7.8 0.69

(From Ginsburg, 1968) M3 15 7.9 53

E. haplodon M1 10.5-11.4 7.8 -

(From Thenius, 1952) M2 12.0-12.8 8.9-9.4 -

M3 15.0-16.4 8.8-9.1 -

E. sansaniensis M1 10.3-10.4 7.0-7.3 -

(From Thenius, 1952) M2 11.1-11.3 8.2-8.4 -

M3 15.0-17.6 7.5-8.8 - 148

Lower M1 12

10

PUPC 08/100* 8 PUPC 08/101*

PUPC 08/114*

6 PUPC 04/24

Length CO-489

PUPC 69/272 ** 4

2

0 0 2 4 6 8 10 12 14 16 Width a

Lower M2 12

10

PUPC 08/100* 8 PUPC 08/101*

PUPC 04/24

6 PUPC 05/11

Length CO-490

PUPC 69/272 ** 4

2

0 0 2 4 6 8 10 12 14 16 Width b

Lower M3 12

10

PUPC 08/100* 8 PUPC 08/101*

PUPC 05/11 6

CO-491 Length

4

2

0 0 5 10 15 20 25 Width c Plot 23a-c: Bivariate scatter graphs showing comparison of the studied specimens with the type specimens

149

W/L Index

0.9

0.8

0.7

0.6 P3 0.5 P4

0.4 M1

M2 0.3 M3 0.2

0.1

0 1 2 3 4 5 6

Plot 24: Width/length indexes of the cheek teeth of Eotragus sp.

Lower P3-M3 PUPC 08/100*

12 PUPC 08/100*

PUPC 08/100*

PUPC 08/100* 10 PUPC 08/100*

PUPC 08/101*

8 PUPC 08/101*

PUPC 08/101*

PUPC 08/114* 6

Length PUPC 04/24

PUPC 04/24

4 PUPC 05/11

PUPC 05/11

2 CO-489 CO-490

PC-GCUF 08/01 0 PUPC 69/272 ** 0 5 10 15 20 25 PUPC 69/272 ** Width

Plot 25: Plot shows the size variation in the lower dentition of Eotragus sp.

150

Lower M1

12

10 E. halamagaiensis E. haplodon 8 E. sansaniensis Eotragus large sp. (PUPC 69/272 ** ) 6 Eotragus sp. (CO-489)

Length Eotragus sp. (PUPC 08/114*)

Eotragus sp. (PUPC 04/24) 4 Eotragus sp. (PUPC 08/101*)

Eotragus sp. (PUPC 08/100*)

2

0 0 2 4 6 8 10 12 14 16 Width a

Lower M2

12

10 Eotragus sp. (PUPC 04/24) Eotragus sp. (PUPC 05/11) 8 Eotragus sp. (CO-490) Eotragus sp. (PUPC 69/272 **) Eotragus sp. (PUPC 08/101*) 6

Eotragus sp. (PUPC 08/100*) Length E. haplodon E. artenensis 4 E. halamagaiensis E. sansaniensis 2

0 0 2 4 6 8 10 12 14 16 Width b

Lower M3

12

10

E. artenensis

E. halamagaiensis 8 E. haplodon

E. sansaniensis 6

Eotragus sp. (PC-GCUF 08/01) Length Eotragus sp. (PUPC 05/11)

4 Eotragus sp. (PUPC 08/101*)

Eotragus sp. (PUPC 08/100*)

2

0 0 5 10 15 20 25 Width c Plot 26a-c: Variation in the dentition of various species of genus Eotragus. 151

Lower M1-M3 E. artenensis E. halamagaiensis 12 E. haplodon E. sansaniensi Eotragus sp. (PC-GCUF 08/01) Eotragus sp. (PUPC 05/11) 10 Eotragus sp. (PUPC 08/101*) Eotragus sp. (PUPC 08/100*) E. halamagaiensis E. haplodon 8 E. sansaniensis Eotragus large sp. (PUPC 69/272 **) Eotragus sp. (CO-489) Eotragus sp. PUPC 08/114* 6 Eotragus sp. (PUPC 04/24)

Eotragus sp. (PUPC 08/101*) Length Eotragus sp. (PUPC 08/100*) Eotragus sp. (PUPC 04/24) 4 Eotragus sp. (PUPC 05/11) Eotragus sp. (CO-490) Eotragus large sp. (PUPC 69/272 **) Eotragus sp. (PUPC 08/101*) 2 Eotragus sp. (PUPC 08/100*) E. haplodon E. artenensis E. halamagaiensis 0 E. sansaniensis 0 5 10 15 20 25 Width

Plot 27: Comparison of the studied specimens with different species of genus Eotragus. 152 Discussion Eotragus is known from the Late Early Miocene of Europe (15 Ma, MN6; Mein, 1989; Steininger et al., 1989; Gentry et al., 1999), Pakistan (18–5 Ma; Solounias et al., 1995; Khan et al., 2008a; Khan, 2009) and from the Middle Miocene of China (16 Ma; Ye, 1989). E. noyei was erected by Solounias et al. (1995) on the basis of a cranial specimen, a horn core (type specimen) and five postcranial specimens recovered from the Lower Siwaliks, with an estimated age between 18.0 and 18.3 Ma (Barry and Flynn, 1989). To date, it is considered as the oldest and smallest species of the genus (Solounias et al., 1995). The oldest representative of the genus is E. artenensis from the middle Burdigalian of Artenay, France, which is a slightly smaller species than others in the genus (Ginsburg and Heintz, 1968). E. haplodon from the Middle Miocene (MN6) locality of Göriach, in the Steirmark Basin of Austria, is relatively larger, has an oval horncore cross-section, horncore abruptly attenuates from base to apex, lower premolars are relatively elongated, and molars are slightly brachydont with a basal pillar. E. sansaniensis from the MN6-MN7 localities in the Sansan Basin of France in addition to Leoben in the Steirmark basin of Austria, is a rather derived species with a size that approaches E. haplodon, horncores are more laterally compressed and gradually attenuate dorsally, dentition is more hypsodont than the former two species, basal pillar is absent, and lower molars are more narrow (Ye, 1989) (Table 10). E. artenesis from Artenay, France, is similar to E. noyei and approximately of the same age (Solounias et al., 1995). E. cristatus from Velheim is approximately 15.5 Ma old. E. haplodon, which is probably conspecific with E. sansaniensis is known from the Austrian locality of Goriach (15 Ma, MN6, Mein, 1989; Steininger et al., 1989). Hamilton (1973) described Eotragus sp. from Gebel Zelten, Libya, which was reported to have an age of 16.5 Ma (Mein, 1989). Tchernov et al. (1987) reported the occurrence of E. cf. sansaniensis in the Negev of Israel. E. halamagaiensis, is known from China. Its age is reported as middle Miocene by Ye (1989). Moya-Sola (1983:pl. 1, figs. 1-2) reported Eotragus sp., from Can Canals and Bunyol (Valencia), and E. cf. artenensis, from Corcoles, Spain. The dentitions of E. artenensis, E. sansaniensis, and from Bunyol have less inclined buccal walls, larger lingual cingula, larger entostyles, and larger molar metaconules than E. sp. The buccal walls are not as aligned as in E. sansaniensis, whereas in E. artenensis they are aligned, forming a straight line (Solounias et al., 1995). Several morphological features, such as the crests of the cusps join up earlier in wear; styles, 153 stylids and ribs are less bulky; obliquely situated hypoconulid, weaker cingula and the confluence of the metaristid, protocristid and postmetacristid are characters corresponding to the genus Eotragus (Gentry, 1999; Rössner, 2006). However, the teeth are too large (Table 10) for E. noyei described from the Kamlial Formation (Solounias et al., 1995), Hasnot, Dhok Pathan formation (Khan, 2007; Khan et al., 2009) and Eotragus sp. (Large size) Dhok Bun Ameer Khatoon (Khan et al., 2008b). The specimens under study are square and tetratuberculate; these can be referred to some herbivorous mammals. Since the cusps are crescentic in outline, so it can be included in sub-order ruminantia of the order artiodactyla. Morphological and metrical features of the specimen clearly indicate a small sized Miocene bovid. Enamel layer is finely rugose, can be referred to family bovidae. The studied cheek teeth differ from cervids in being higher; the crests of the cusps join up earlier in wear; styles, stylids, and ribs are less bulky and cingula are weaker (Gentry, 1999). The studied specimens (PUPC 08/100, PUPC 08/101, and PUPC 08/114) are clearly comparable with specimens recovered by Solounias et al. (1995), and Khan et al. (2008b) from the Lower Siwaliks. PUPC 08/114 compare positively with first molar of PUPC 69/272 large sp. (Table 10). The size variation is observed among different species of the genus Eotragus (Table 10 & 11, Plots 23a-c, 24, 25 and 26). Plot 27 provides comparison of the studied specimens with different species of genus Eotragus. Therefore, the material is referred to Eotragus sp. but more material is needed for precise species identification.

154 DISCUSSION OF THE EVEN AND ODD-TOED MAMMALS

The Siwalik formations of northern Pakistan consist of deposits of ancient rivers that existed throughout the early Miocene through the late Pliocene. The formations are highly fossiliferous with a diverse array of terrestrial and freshwater vertebrates, which in combination with exceptional lateral exposure and good chronostratigraphic control allows a more detailed and temporally resolved study of the sediments and faunas than is typical in terrestrial deposits. Consequently the Siwaliks provide an opportunity to document temporal differences in species richness, turnover, and ecological structure in a terrestrial setting, and to investigate how such differences are related to changes in the fluvial system, vegetation, and climate (Willis and Behrensmeyer, 1994; Johnson et al., 1985). It is quite possible that the progressive development of the Siwalik faunas was dependant upon a change from the relative dry flood plain environment to a moister floodplain and forest environment. The Himalayan mountain mass was comparatively low, and therefore less effective as a trap for moister laden winds than are the present mountains at the beginning of the Siwalik deposition. Consequently the rising of the Himalaya may have caused a progressive increase in moister and in temperature in the northern portion of India, due to the fact that the successively higher mountains caused an increase in precipitation along their southern flanks. Thus the Himalayas have acted as an increasing effective barrier in isolating India from the rest of Asia, cutting out the cold temperatures sweeping down from the north and the east and retaining the warm moister laden winds blowing in from the Indian Ocean (Cerling, et al., 1997; Johnson et al., 1982; Barry et al., 2002). The climate changes in the Neogene were of low-amplitude in a generally warm-humid background. Those of the Quaternary were of higher amplitude with a longer phase. Cool or cold dry and slightly warm semiarid and semi-moist climates alternated on a generally cool-dry background (Ojha et al., 2000). Lithofacies of the Chinji Formation are thought to represent deposits of either the Paleo-Indus River or a similar axial fluvial system (Johnson et al., 1982). The multistoried channel type sandstone-bodies of Chinji Formation suggest a consistent flow direction to the SSE (Abbasi, 1988). There are broad similarities between channel geometrics, discharge and sedimentary characters of Siwalik rivers and modern Indus river system including emergence from mountain belt, generally parallel flow to the basin axis, slopes range and bankfull discharge(Mackey and Bridge, 1995). 155 In spite of adequate exposure of Chinji Formation in the DBAK, fossils are not easily found in this area as compared to Chinji stratotype. It is particularly because of steeper dips and some structural complication, thus reducing the area of exposure per bed and also an increase in mudstone indicates existence of floodplains, which relatively have less potential for fossilization. The Giraffokeryx remains in this study document the vegetation of Chinji Formation as seasonal woodland with riparian areas of forest. The occurrence of Giraffokeryx punjabiensis from Chinji–Nagri Formation testifies a change in the composition of the Siwaliks from seasonal woodland – tree savannas. Giraffokeryx Punjabiensis was a mixed feeder and is well adapted to a life in savannas. (Franz- Odendaal and Solounias 2004; Janis and Scott, 1987a; Solounias et al., 2000; Solounias and Moelleken, 1993). The Siwalik vertebrate fauna including medium or large mammals that might have been exclusively forest species. There were also few species that appear to represent adaptations to open grassland. Prehipparion fauna contain more primitive elements but these are not necessarily forest species. Less common taxa including tragulids and Chalicotheres are either absolutely less abundant or had different more restricted patterns. In general the nature of the fauna in this basin suggests the occurrence of open grassland and savannas in the late Middle Siwaliks. The pattern of species disappearance and appearance suggests that biotic interactions may have been more important than environmental change. The 7.8 Ma events are characterized solely by appearances and that at 7.3 Ma by a combination of appearances and disappearances. These two latest Miocene events include more taxa that were shorter ranging and less common, a difference of mode that developed between approximately 9.0 and 8.5 Ma. Both events also show a close temporal correlation to changes in floodplain deposition and vegetation. The 7.8 Ma event follows the widespread appearance of C4, vegetation and is coincident with the shift from equid- dominated to more evenly balanced large-mammal assemblages. The 7.3 to 7.0 Ma event starts with the first occurrence of C4 -dominated floras and ends with the last occurrence of C3- dominated vegetation. Absence of a consistent relationship between depositional facies and the composition of faunal assemblages leads us to reject fluvial system dynamics as a major cause of faunal change (Barry et al., 2002). The 10.3 Ma event marks the extinction of the Giraffokeryx-Listriodon fauna, which historically has been linked to the first appearance of equids in the Siwaliks. At 10.3 Giraffokeryx punjabiensis is replaced by Bramatherium megacephalum. A marked 156 size increase also occurs between 11 and 10.3 Ma with the first appearance of the Selenoportax and Tragoceridus. Both giraffes and bovids exhibit continuing size increases after 10 Ma. The 7.8 Ma turnover is also coincident with the shift from equid-dominated to more evenly balanced large-mammal assemblages. Appearing taxa include a hypsodont rodent and a high-crowned tragulid, implying a grass and possibly C4 diet. The carbon isotope event indicates that habitats with C4, plants became more common on the floodplain. This is strong evidence of changing habitat availability and is concordant with the appearance of grazing or mixed-feeding taxa. At the same time, the paleosol carbon record indicates that C3-dominated habitats still persisted and were even common, thus explaining the absence of elevated extinction. The younger turnover event between 7.3 and 7.0 Ma is composed of both appearances and disappearances. It begins very shortly after the first occurrence of C4-dominated floras at 7.37 Ma and ends with the last occurrence of C3-dominated vegetation at 7.04 Ma. These two isotopic thresholds are critical to understanding the impact of this vegetation change on the mammals, in that they record the appearance and disappearance of end-members from the vegetation spectrum. They thus indicate both the gain and the loss of types of extreme open and closed habitats, and are likely related directly to the appearance and disappearance of individual species. This same broad interval also gives evidence of increasingly seasonal stream flow and perhaps higher levels of habitat disturbance. The change in large- mammal diversity between 10 and 9.8 Ma potentially could be related to fluvial system dynamics, as it closely follows the initial appearance at 10.1 Ma of the interfan Dhok Pathan system. The nature of the fauna in this basin suggests the occurrence of open grassland and Savannahs in the late Middle Siwaliks.

157

Age and Correlation The even and odd-toed mammals identified from the Dhoke Bun Ameer Khatoon on the basis of the published and unpublished work by Cheema, 2003, Khan et al., 2008a and the present study is as follows:

Family Suidae Listriodon pentapotamiae Conohyus sindiensis Family rhinocerotidae Gaindatherium browni Brachypotherium perimense Chilotherium intermedium Family Chalicotheriidae Chalicotherium intermedium Family Bovidae Tribe Boselaphini Sivaceros gradiens

Eotragus noyei Eotragus sp. Elachistocerus sp. Tribe Antilopini Gazella sp. Gazella lydekkeri Family Tragulidae Dorcatherium majus Dorcatherium minus

Family Giraffidae Giraffa priscilla Giraffokeryx punjabiensis

The age of the even and odd-toed mammals identified from the Dhoke Bun Ameer Khatoon is clearly later than the early Miocene, and also they appear to be earlier than the Pliocene, because boselaphines are part of the even and odd-toed fauna, likely to be the end representatives of the late Miocene. Even and odd-toed mammals found in the Dhoke Bun Ameer Khatoon belong to the families Bovidae, Tragulidae, Suidae, Rhinocerotidae and Giraffidae. Thus, this makes the recovered even and odd-toed mammals fauna from DBAK respectable enough to provide an indication for the age. Whereas giraffids, which are present very common in Village DBAK. The suid (10.2-6.5 Ma) and the antilopini 158 (8.6-7.4 Ma) suggest an age of 7-5 Ma for DBAK. In the present work, all the collected specimens come from the lower Siwaliks. Several descriptive phases exist for the late Miocene (Vallesian and mainly Turolian) faunas that are known from Greece, through Turkey to Iran, known as the Graeco-Iranian province (Iliopoulos, 2003). Knowledge of these faunas originated from the discovery and study of the major localities of Pikermi (Greece), Samos (Greece), and Maragheh (Iran), known since the 19th century (Gentry, 1999; Solounias, 1982a). Gazella are considered as typical late Miocene taxa. Hence, a date around 7.0 Ma or during the latest Miocene (Barry et al., 1991 for dating of the Siwalik deposits) would be considered as a possible date for a fauna containing. Based on the fossil record, it seems that Sivatheres do not appear before the late Miocene, unless one adopts the doubtful proposition that the four horns of the middle Miocene Giraffokeryx indicate a sivatheriine affinity (Gentry, 1999). G. punjabiensis comes from the Dhok Pathan (Colbert, 1935) in the time span between 7.1-5.0 Ma (Barry et al., 1991). G. punjabiensis is abundant in the DBAK localities, having a size similar to that of Palaetragus coelophrys from Maragheh (Iran). The tragulids (Dorcatherium) that appear in the Siwalik sequences are dated as 18- 6.4 Ma (Barry and Flynn, 1989). The genus Dorcatherium is close to the genus Saimotragulus, found in the Middle Miocene of Thailand and the genus Yunnanotherium, reported from the late Miocene of China (Vislobokova, 2001). Therefore, the even and odd-toed mammals of DBAK support a late Miocene perhaps equivalent to MN 13 (late Turolian to late Vellasian) in terms of European Mammal Neogene zone scale and around 7.0-5.0 Ma in terms of dating. The DBAK localities can be compared with those late Miocene faunas in adjacent continental regions. Making comparisons at regional level, rather than with particular localities, increases the deficiency of temporal regions. Below tribal level the DBAK even and odd-toed mammals show little resemblance to the rich Turolian assemblage. Gazella lydekkeri, the much greater abundance of spiral-horned Antilopini is also noteworthy in the Graeco-Iranian Turolian province. The following list of even and odd-toed mammals for the Graeco-Iranian region is based mainly on Solounias (1981), Bernor (1986), Kohler (1987), and Gentry and Heizmann (1996).

159 Family Giraffidae Palaeotragus, 2 spp. (P. rouenii and P. coelophrys) Samotherium boissieri and S. major Decennatherium macedoniae Helladotherium duvernoyi and H. grande Bohlinia attica Family Bovidae Tribe Boselaphini Tragoportax sp. Tribe ?Bovini Samokeros minotaurus Tribe Reduncini ?Redunca sp. Tribe Antilopini Prostrepsiceros sp. or P. vinayaki Protragelaphus sp. Nisidorcas planicornis (?Hispanodorcas) rodleri Oioceros sp. Samodorcas kuhlmanni Gazella sp. (G. capricornis) Tribe Ovibovini Urmiatherium sp. Plesiaddax inundatus Criotherium argalioides Palaeoryx pallasi Tribe Caprini Protoryx carolinae Pachytragus sp. Norbertia hellenica Pseudotragus sp.

Bovini, Boselaphini and Antilopini are present in the Turolian of the Graeco-Iranian province as well as in the DBAK. There are rare records of Tragulidae in the Graeco- Iranian Turolian.

In Langebaanweg (South Africa) the following species are present (Harris, 1976; Gentry, 1978).

160 Family Giraffidae Giraffa sp. Sivatherium hendeyi Family Bovidae Tribe Tragelaphini Tragelaphus ?sp. Tribe Boselaphini Tragoportax acrae Tribe Bovini Simatherium demissum Tribe Reduncini Kobus sp. Tribe Alcelaphini Damalacra sp. Tribe Neotragini Raphicerus paralius Tribe Antilopini Tribe Ovibovini Gen. et sp. Indet.

Most of the species show primitive characters and are thereby differentiated from later species of East and South Africa. Only Giraffa sp. and Gazella sp. has a resemblance to a species from the DBAK and this is likely to reflect that both species are late Miocene. Gentry (1999) identified the following species from the Late Miocene Baynunah Formation (Abu Dhabi):

FAMILY GIRAFFIDAE ?PALAEOTRAGUS ?Bramatherium sp. Giraffidae sp. Indet. Family Bovidae Tribe Boselaphini Tragoportax cyrenaicus Pachyportax latidens Tribe indet. Bovidae sp. Indet Tribe Antilopini Prostrepsiceros aff. libycus Prostrepsiceros aff. vinayaki Gazella aff. lydekkeri

Tragulidae are absent in the Baynunah Formation, however some Bovidae and Giraffidae species are common with the localities of the DBAK. In East Africa, at Mpesida, Lukeino, and the Manonga Valley (Thomas, 1984; Gentry, 1997), the combined giraffid and bovid fauna can be given as: 161

Family Giraffidae Giraffa sp. Sivatherium sp. Family Bovidae Tribe Tragelaphini Tragelaphus sp. Tribe Bovini Ugandax cf. gautieri Tribe Cephalophini Cephalophus sp. Tribe Reduncini Kobus aff. subdolus Kobus aff. porrecticornis Tribe Hippotragini Praedamalis sp. Tribe Alcelaphini Damalacra sp. Alcelaphini, sp. Tribe Aepycerotini Aepyceros sp. Tribe Neotragini Madoqua sp. Tribe Antilopini ?Gazella sp.

This assemblage has some resemblance to Langebaanweg (South Africa), but Cephalophini, Hippotragini, and Aepyceros are additional African constituents. The only resemblance to the DBAK even and odd-toed mammals seems to be the presence of a giraffids (Giraffa sp.) and bovids (Gazella sp.) at a comparable stage of evolutionary development. There are no Boselaphini or spiral horned Antilopini, although boselaphines were present earlier in the East African Miocene at Lothagam 1 (Kenya). Also, Nakaya et al. (1984: 109) recorded a possible spiral horned antilopini (as Palaeoreas sp.) in the earlier Namurungule Formation, and Smart (1976) listed Antilope sp. for Lothagam 1 (Kenya). The North African pecorans, as known from Bled ed-Douarah, Sahabi, and Wadi Natrun (Saudi Arabia) (Geraads, 1989; Harris, 1987b; Lehmann and Thomas, 1987), are:

Family Giraffidae Giraffidae, sp. indet. Sivatherium aff. hendeyi Family Bovidae Tribe Boselaphini Tragoportax cyrenaicus Tribe Bovini “Leptobos” syrticus Tribe Reduncini Kobus sp. 162 Tribe Hippotragini Hippotragus sp. Tribe Alcelaphini Damalacra sp. Tribe Neotragini Raphicerus sp. Tribe Antilopini Prostrepsiceros libycus Gazella sp.

The African tribes in the list are very close to the DBAK even and odd-toed mammals with a conspecific spiral-horned antilopini and giraffids, but they differ in the presence of several exclusively African tribes. It needs to be remembered that several stratigraphic levels are represented at these localities, probably ranging up into the Pliocene (Geraads, 1989) and some of the African tribes of antelopes may be present only in the higher levels.

The DBAK even and odd-toed mammals are different from those of the Graeco- Iranean Turolian province. Where genera are common, the species are different, and the DBAK ones look earlier. They also differ from East and South African faunas in that the giraffid sp. indet. is not a Giraffa, and in the absence of African tribes of antelopes. The DBAK fauna shows some affinity with the Baynunah Formation (Abu Dhabi) and the North African faunas. While deer are common in late Miocene faunas north of the present Black Sea, they are rare in Greece, Turkey, Iran, and the Late Miocene of the Siwaliks, and until now they are absent from Abu Dhabi, and North Africa. The presence of giraffines, boselaphines, antilopines, and a gazelle at Abu Dhabi, North Africa and the Siwaliks could indicate a rather parkland-like landscape. The presence of a southern or hot climate is plausible, however, without any substantial development of aridity. The gazelle might have had a life style more like that of the present day West African Gazella rufifrons than the lifestyle of the desert gazelles of North Africa and Arabia. The presence of the tragulids in the Late Miocene of the Siwaliks and their absence from North Africa and Abu Dhabi suggests that the range contraction of this family had proceeded further, and that suitable habitats were unavailable in the Arabian Peninsula.

The presence of Chilotherium, Dorcatherium sp., Eotragus sp., Eotragus large size sp. and a listriodont suid in the lower stratum suggest and Early Miocene age (Welcomme et al., 2001, Antoine and Welcome, 2000). Eotragus is known from the late early Miocene of Europe (Gentry et al., 1999), Pakistan (Solounis et al., 1995) and from the middle Miocene of China (Ye, 1989) but the presence of this taxon in the DBAK site considerable extends its geographical distribution. 163 From Pilgrim (1912) to onward several authors believed that the Bugti fauna is Miocene in age, but revision of previously collected specimens have led to propose new interpretation of biostratigraphy of the Bugti hills (Metais et al., 2003). For the comparison of various faunal assemblages and quantity similarities and differences, Simpson (1945, 1960) method is considered best among all others to compare various faunal assemblages. The mammalian fossil fauna of North Africa resembles with East Africa as compare to southern Europe, whereas there is not any resemblance of any genera between Spanish and Siwalik assemblages. While the Siwaliks assemblage shares more taxa with East African sites. African assemblages in general, show more similarities with the Spanish assemblage than that of the Siwalik assemblages (Hail-Sellassie et al., 2003). The Siwalik fauna can be clearly separated from other localities under comparison and this is more likely to be a consequence of either geographic remoteness of presence of different barriers for possible faunal interchange during these times. In general, the quantitative results indicate a higher faunal interchange within Africa during the Late Miocene than between Africa and either Europe or Asia. However, it should also be noted that the presence of taxa shared exclusively between these regions. However, immigration is one of the most powerful processes that affect diversity of a given faunal assemblage. Miocene genera are common in Europe, Africa and India. During much of the Tertiary Africa and India were isolated. Major difficulties are more likely to be assigned to a new species, than to be subsumed with a conspecific species geographically farther away (Colbert, 1935, Beck and Burbank, 1990, Barry and Flynn, 1990, Hail-Selassie et al., 2003). From above review it is clear that much more work is required to correct such biases. Conclusion All the described specimens are collected from the Chinji Formation because the Chinji Formation is well exposed in the study area, although the Nagri and Dhok Pathan Formation are also exposed but author could not collect any specimen during the particular field work from these Formations. The base of Dhok Pathan Formation is only exposed in the study area and perhaps the structural complication is one of the basic reason that the fossils are very rare in the study area.

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APPENDIX List of Studied Specimens and their Dental Position.

SERIAL NUMBER DENTAL POSITION Family Rhinocerotidae Gaindatherium browni

PUPC 08/14 Right P2

PUPC 08/85 Left mandibular ramus with M2-3 Family Suidae Listriodon pentapotamiae

PUPC 08/9 Right mandibular fragment with P4-M3

PUPC 08/9 Left mandibular fragment with M1-3 PUPC 08/9 Left side upper incisor

PUPC 08/21 Right P4 PUPC 08/24 Right upper canine

PUPC 08/70 Left M1

PUPC 08/93 Left P 3 Family Tragulidae Dorcatherium majus PUPC 08/23 Right maxillary fragment with M2-3 PUPC 08/36 Right M2 PUPC 08/42 Left M2 Dorcatherium minus PUPC 08/90 Left M1 Family Giraffidae Giraffa priscilla

PUPC 08/10 Left mandibular ramus with M1-2

PUPC 08/29 Right M2 192 Giraffokeryx punjabiensis

PUPC 08/11 Left M3

PUPC 08/13 Right M2 PUPC 08/18 Right M2 PUPC 08/26 Left P4 PUPC 08/28 Right M2 PUPC 08/31 Right M1 PUPC 08/33 Right P3 PUPC 08/34 Left P3 PUPC 08/35 Left P2

PUPC08/38 Right P3 PUPC 08/43 Left P3

PUPC 08/45 Left M1

PUPC 08/94 Left P3

PUPC 08/95 Left P3 PUPC 08/96 Right P4 PUPC 08/97 Right P2 PUPC 08/98 Left P3 PUPC 08/99 Left M2 PUPC 08/105 Right M1 PUPC 08/113 Left M1 PUPC 08/117 Right M3 Family Bovidae Gazella sp. PUPC 08/16 Right mandibular ramus having M1-2 PUPC 08/111 Left M2

PUPC 08/116 Left mandibular ramus having M1-2. Eotragus sp.

PUPC 08/100 Right mandibular ramus having P3-M3.

PUPC 08/101 Left mandibular ramus having M1-3

PUPC 08/114 Right M1