TAXONOMY AND DENTAL MORPHOLOGICAL STUDIES OF FOSSIL ARTIODACTYLS FROM TYPE LOCALITY OF DHOK PATHAN, PUNJAB, PAKISTAN
MAHBOOB IQBAL
DEPARTMENT OF ZOOLOGY UNIVERSITY OF THE PUNJAB LAHORE, PAKISTAN
2010
TAXONOMY AND DENTAL MORPHOLOGICAL STUDIES OF FOSSIL ARTIODACTYLS FROM TYPE LOCALITY OF DHOK PATHAN, PUNJAB, PAKISTAN
By
MAHBOOB IQBAL
A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF
Doctor of Philosophy
In the Faculty of Life Sciences, University of the Punjab, Lahore, Pakistan
Supervisor
Prof. Dr. Muhammad Akhtar
Co-Supervisor
Dr. Muhammad Akbar Khan
DEPARTMENT OF ZOOLOGY UNIVERSITY OF THE PUNJAB LAHORE, PAKISTAN
August, 2010
DEDICATION
DEDICATED WITH LOVE TO
MY PARENTS MY FAMILY (DR. SAMINA, UMAR & HADEEQA) AND ALL THOSE WHO LOVE, CARE AND PRAY FOR ME.
ABSTRACT
A taxonomic investigation of Late Miocene artiodactyl fauna of Dhok Pathan, the Middle
Siwaliks, Pakistan, was undertaken. Comparative morphometric features of the late
Miocene artiodactyls from the locality are studied in this thesis. The late Miocene artiodactyls are presented on the basis of description of abundant material from the Dhok
Pathan type locality of the Dhok Pathan Formation, the Middle Siwaliks and the remains increasingly indicate taxonomic diversity. Quantitatively, the taxa of bovids are the most predominant. But cervids, tragulids, giraffids and suids are approximately as common as each other at Dhok Pathan.
More than 130 artiodactyl fossil specimens were studied in this thesis from the Dhok
Pathan type locality. All the described specimens include Skulls, horn cores, isolated teeth and fragments of maxillae or mandibles. These fossils document twenty three artiodactyl species belonging to fifteen genera and five families. This assemblage includes two new taxa: Miotragocerus large sp. and ?Hydaspitherium sp. Boselaphines, antilopines, reduncines, cervids, tragulids, giraffids and suids are abundant in the locality whereas tragelaphines and alcelaphines are absent.
Biogeographically, the late Miocene artiodactyls indicate strong relationships with
Eurasian and African late Miocene sites. In this study, paleoenvironmental interpretations of the type locality provide important evidence regarding late Miocene paleoenvironments. Most of these taxa indicate a predominance of woodland to savannah habitat during the deposition of the Dhok Pathan Formation.
ACKNOWLEDGEMENTS
All praises with my deepest gratitude to Almighty Allah, whose uniqueness, oneness and wholeness in unchallengeable. All and every respect is for His Holy Prophet HAZRAT MOHAMMAD (Peace Be Upon Him), who enlightened us to understand the mystery of life. I acknowledge the depth of my gratitude and heart full thanks to highly learned and respected project Design Supervisor professor Dr. Mohammad Akhtar, Department of Zoology university of the Punjab under whose auspices I took a stride in completion of this work. He is also thanked for granting permission to use some bovids, suids, material collected by him and housed in the Dr. Abu Bakr Fossil Display and Research Centre and Paleontology laboratory of the university of the Punjab for this study. I feel great pleasure and honour to express my deep sense of gratitude to my respected and genius Co Supervisor Dr. Mohammad Akbar Khan assistant Professor Zoology GC University, Faisalabad for their constructive suggestions, guidance and encouragement during my research project. I feel highly privileged in taking the opportunity to thank Professor Dr. Asif Mehmood Qureshi, Prof. Mirza Afzal Baig and Prof. Abdul Qayyum Khan of Zoology Department Govt. College of Science Wahdat Road Lahore, for their moral support and encouragement and interest during the course of this work. I owe special thanks to Dr. Abdul majid Khan, Dr. Umar Farooq and Dr. Abdul Ghaffar who supported me at every critical stage during the course of my research project. I am also thankful to Mr. Abdul Rauf Malik Associate Prof. of Zoology Govt. College Talagang (Chakwal) providing Base Camp for the fossil Collection. I sincerely thank to Mr. sajjid Shah, Mr. Maqsood Ahmed and Mr. Abdul Razzaq for joining me in field trips, for the fossil collection and for serving me in the laboratory. Mr. Adeeb Babar is thanked for his help with the preparation for the maps and photographs.
MAHBOOB IQBAL Dated: August, 2010 i
LIST OF ABBREVIATIONS
Ca Circa Myr Million years Ma Million years ago MN European Mammal Neogene zone scale GPTS Geomagnetic Polarity Time Scale GRTS Geomagnetic Reversal Time Scale DP1-DP17 Localities of Dhok Pathan from numbers1 to 17 AMNH American Museum of Natural History BMNH British Museum of Natural History PMNH Pakistan Museum of Natural History PUPC logy, Punjab University, Lahore, Pakistan.
PC-GCUF Palaeontological Collection of GC University, Faisalabad, Punjab, Pakistan.
P or p Upper or lower premolar
M or m upper or lower molar GSI Geological survey of India GSP Geological Survey of Pakistan DP Deciduous Premolar W/L Width/Length ratio Fms/Fm Formations/Formation. r Right l Left mm Millimeters
ii
TABLE OF CONTENTS
Abstract
Acknowledgements
List of Abbreviations
Table of Contents
List of Tables
List of Figures
Chapter 1
Introduction
Geography
Geology
Stratigraphy and Chronology
Biostratigraphy
Siwalik Artiodactyls
Objectives of Present Study
Thesis Layout
Chapter 2
Materials and Methods iii
Taxonomic Description
Chapter 3 Systematic Palaeontology Family Bovidae Tragoportax Tragoportax punjabicus Tragoportax browni Tragoportax cf. salmontanus Miotragocerus Miotragocerus large sp.
Miotragocerus cf. gluten Pachyportax Pachyportax latidens Pachyportax cf. nagrii
Selenoportax Selenoportax cf. vexillarius Prostrepsiceros Prostrepsiceros cf. vinayaki Gazella Gazella lydekkeri Kobus Kobus porrecticornis cf. Kobikeryx sp. Caprotragoides Caprotragoides cf. potwaricus Family Cervidae iv
Cervus Cervus cf. sivalensis Cervus cf. triplidens Family Tragulidae Dorcatherium Dorcatherium majus Dorcabune Dorcabune anthracotherioides Dorcabune cf. nagrii Family Giraffidae Hydaspitherium
Hydaspitherium cf. megacephalum
Hydaspitherium cf. grande
?Hydaspitherium sp.
Family Suidae
Propotamochoerus
Propotamochoerus hysudricus
Hippopotamodon
?Hippopotamodon sp.
Chapter 4 Palaeoenvironment and Paleobiogeography of the Dhok Pathan Artiodactyls
Chapter 5 Conclusions
References
v
Appendices
LIST OF TABLES
Table 1: Mammalian faunas of Dhok Pathan. Many species are under taxaonomic revision. Table 2: Comparison of the cranial measurements (mm) of the Siwalik Tragoportax. Table 3: Comparative measurements of the cheek teeth of T. punjabicus in mm. Table 4: Comparative measurements of the cheek teeth of T. browni Pilgrim in mm. Table 5: Comparative measurements of the cheek teeth of Miotragocerus sp. and Miotragocerus gluten (Pilgrim, 1937) in mm. Table 6: Comparative measurements of the cheek teeth of P. latidens (Lydekker) Pilgrim in mm. Table 7: Comparative measurements of the cheek teeth of S. cf. vexillarius in mm. Table 8: Comparative measurements (mm) of the cheek teeth of G. lydekkeri. Table 9: Comparative measurements of the cheek teeth of the Siwalik Cervus in mm. Table 10: Comparative measurements of the cheek teeth of D. majus and D. minus in mm (millimeters). vi
Table 11: Comparative measurements of the cheek teeth of Dorcabune anthracotherioides and D. nagrii in mm. Table 12: Comparative measurements of the lower cheek teeth of the giraffids in the Dhok Pathan type locality. Table 13: Comparative measurements of the cheek teeth of Propotamochoerus hysudricus and ?Hippopotamodon sp. in mm (millimeters).
APPENDICES Appendix 1: List of the studied artiodactyls from Dhok Pathan. Appendix 2: Studied material.
Appendix 3: The published article.
LIST OF FIGURES
Figure 1. A. Location of the Potwar Plateau in northern Pakistan; the studied areas are encircled. B. Simplified geology map of the Dhok Pathan area indicating the fossiliferous localities around the Dhok Pathan Rest House (DP – abbreviation for Dhok Pathan) from where the studied material is recovered. Figure 2. Percentage of species represented in the Dhok Pathan mammalian fauna. Figure 3. Tragoportax punjabicus. 1. PUPC 86/149 opisthocranium: a) dorsal view, b) ventral view, c) occipital view. 2. PUPC 86/150 opisthocranium: ventral view. Scale bar equals 30 mm total. Figure 4. Tragoportax punjabicus. 3. PUPC 86/264 left horn core: a) lateral view, b) medial view. Scale bar equals 30 mm total. Figure 5. Tragoportax punjabicus. 4. PUPC 08/126 palatine parts of both maxillae with right P3-M3 and left P2-M3: occlusal view. 5. PUPC 83/275 right maxillary fragment with P3-M3. 6. PUPC 97/12 left mandible fragment with p2-p4. 7. PUPC 83/277 left mandible vii
fragment with m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 6. Tragoportax browni. 1. PUPC 86/204 opisthocranium: a) dorsal view, b) ventral view, c) occipital view. Scale bar equals 30 mm total. Figure 7. Tragoportax browni. 2. PC-GCUF 09/11 horn core, side indeterminate: a) anterior view, b) posterior view, c) medial view, d) lateral view. Scale bar equals 30 mm total. Figure 8. Tragoportax browni. 3. PUPC 86/151 left maxillary fragment with M1-M2. 4. PUPC 83/265 left mandible fragment with m2-m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 9. Tragoportax cf. salmontanus. 1. PUPC 93/20 left horn core: a) anterior view, b) posterior view, c) medial view. Scale bar equals 30 mm total. Figure 10. Scatter diagram showing horn-cores and dental proportions of the Dhok Pathan Tragoportax. Figure 11. Miotragocerus large sp. 1. PUPC 83/209 left maxillary ramus with P2-M1. 2. PUPC 87/246 left maxillary ramus with DP3-M2. 3. PUPC 09/86 left mandibular ramus with p4-m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 12. Miotragocerus cf. gluten. 1. PUPC 01/01 left horn core: a) anterior view, b) medial view, c) lateral view, d) posterior view. Scale bar equals 30 mm total. Figure 13. Scatter diagram showing dental proportions of the Dhok Patan Miotragocerus. Figure 14. Pachyportax latidens. 1. PUPC 96/18 right horn core: a) anterior view, b) medial view, c) posterior view, d) lateral view. Scale bar equals 30 mm total. Figure 15. Pachyportax latidens. 2. PUPC 80/646 isolated M2, side indeterminate. 3. PUPC 09/74 isolated M2, side indeterminate. 4. PUPC 80/203 isolated right M2. 5. PUPC 08/87 right mandibular viii
ramus with m3 with broken hypoconulid and an anterior part of m2. 6. Pachyportax cf. nagrii. PUPC 99/50 isolated molar, side indeterminate. Occlusal views. Scale bar equals 30 mm total. Figure 16. Scatter diagram showing dental proportions of the Dhok Pathan Pachyportax. Figure 17. Selenoportax cf. vexillarius. 1. PUPC 99/47 right mandibular ramus with m2 and roots of m1. 2. PUPC 97/02 isolated right m3. 3. PUPC 97/01, isolated right m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 18. Scatter diagram showing dental proportions of the Dhok Pathan Selenoportax. Figure 19. Prostrepsiceros cf. vinayaki. 1. PUPC 87/319 left horn-core. 2. PUPC 87/318 ?right horn-core. a) anterior view, b) medial view, c) lateral view, d) posterior view. Scale bar equals 30 mm total. Figure 20. Gazella lydekkeri. 1. PUPC 83/319 left horn-core. 2. PC-GCUF 09/45 left horn-core. a) anterior view, b) medial view, c) lateral view, d) posterior view. 3. PUPC 08/136 right isolated m2: occlusal view. 4. PUPC 07/13 isolated left m3. 5. PUPC 09/76 isolated left m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 21. Scatter diagram showing dental proportions of Gazella. Figure 22. Kobus porrecticornis. 1. PUPC 87/124A left horn-core. 2. PUPC 87/124B right horn-core. 3. PUPC 02/126 left horn-core. 4. PUPC 87/180 left horn-core. a) anterior view, b) medial view, c) lateral view, d) posterior view. Scale bar equals 30 mm total. Figure 23. cf. Kobikeryx sp. 1. PUPC 99/49 horn core, side indeterminate: a) anterior view, b) posterior view. Scale bar equals 30 mm total. Figure 24. Caprotragoides cf. potwaricus. 1. PC-GCUF 09/44 right horn-core: a) anterior view, b) medial view, c) lateral view, d) posterior view. Scale bar equals 30 mm total. ix
Figure 25. Cervus cf. sivalensis. 1. PUPC 04/18 right mandibular ramus with m2-3: a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 26. Cervus cf. triplidens. 1. PUPC 09/71 right maxillary ramus with P4- M2: a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 27. Scatter diagram showing dental proportions of the Siwalik cervids. Figure 28. Dorcatherium majus. 1. PUPC 03/14 isolated left M2. 2. PUPC 01/21 isolated right m1. 3. PUPC 07/09 isolated left m2. 4. PUPC 01/26 left mandibular ramus with m2 and posterior portion of m1. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 29. Scatter diagram showing dental proportions of the Siwalik Dorcatherium. Figure 30. Dorcabune anthracotherioides. 1. PUPC 99/338 isolated right M2. 2. PUPC 04/01 right mandibular ramus with m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 31. Scatter diagram showing dental proportions of the Siwalik Dorcabune. Figure 32. Dorcabune cf. nagrii. 1. PUPC 96/45 isolated left probably M1: a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 33. Hydaspitherium cf. megacephalum. 1. PUPC 09/78 left mandibular ramus with broken m1, complete m2 and partially erupted m3: a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 34. Hydaspitherium cf. grande. 1. PUPC 09/77 right mandibular ramus with m2-3: a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. x
Figure 35. ?Hydaspitherium sp. 1. PUPC 83/267 right mandibular ramus with m2-3: a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 36. Scatter diagram showing dental proportions of the studied giraffids. Figure 37. Propotamochoerus hysudricus. 1. PUPC 07/32 left mandibular ramus with p4-m3. 2. PUPC 94/65 right mandibular ramus with p4- m2. 3. PUPC 99/02 left mandibular ramus with p4-m1. 4. PUPC 97/15 right isolated m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. Figure 38. ?Hippopotamodon sp. 1. PUPC 00/14 left mandibular ramus with broken p4 and partially complete m1: a) occlusal view, b) lingual view, c) labial view. 2. PUPC 08/26 badly crushed right mandibular ramus with damaged m1-3: occlusal view. Scale bar equals 30 mm total. Figure 39. Scatter diagram showing dental proportions of the studied Dhok Pathan Suoids. Referred data are taken from Colbert (1935) and Pickford (1988). Figure 40. Percentage of species represented in the Dhok Pathan artiodactyls. 1
INTRODUCTION
Geography
The village Dhok Pathan (Lat. 33° 07'N: Long. 72° 14'E) is situated in the district
Chakwal of northern Pakistan (Fig. 1A). The type locality is situated at Soan River about
75 km from Rawalpindi on Rawalpindi-Sargodha road and it is surrounded by extensive
Neogene freshwater sedimentary rocks. The Dhok Pathan fossiliferous area is situated in the vicinity of the Dhok Pathan Rest House near the Soan River at an altitude of 1073 feet (Fig. 1B). The average thickness of the sequence around this area is about 497 m
(Barry et al., 2002). Sites surrounding the village present an abundance of vertebrate fossils that represent almost all major Eutherian groups. The region of the Dhok Pathan exposes the most complete sequence of the Siwalik Group and yields a diversified assemblage of the Middle Siwalik faunas (Table 1). The Dhok Pathan bed near the Dhok
Kundrali is one of the most fossiliferous beds of the Siwaliks (Akhtar, 1992) and most of the material described in this thesis is recovered from the Kundrali unit (DP9-13). This bed has yielded a large number of artiodactyl samples but these are not homogeneous.
This is particularly important because much of the Dhok Pathan fauna of Pilgrim, Brown, and Lewis apparently came from the vicinity of the bed (Barry et al., 2002). It is situated in the vicinity of the Dhok Pathan Rest House (Lat. 33° 07'N: Long. 72° 22'E) which is placed about at the bank of the Soan River near the village Dhok Pathan. Four new localities have been excavated and they are DP8, DP9, DP16 and DP17 (Fig. 1B). 2
Figure 1. A. Location of the Potwar Plateau in northern Pakistan; the studied areas are encircled (map is modified from Behrensmeyer and Barry, 2005 and the boundary dates are from Dennell et al., 2008 and Nanda, 2008). B. Simplified geology map of the Dhok Pathan area indicating the fossiliferous localities around the Dhok Pathan Rest House (DP – abbreviation for Dhok Pathan) from where the studied material is recovered (map is modified from Colbert, 1935). 3
Geology
The geology of the Siwaliks has been well studied and documented by national
international researchers since the 18th century. The British and American workers did the
pioneering work on the Siwalik geology and palaeontology (Lydekker, 1878; Colbert,
1935; Behrensmeyer and Tauxe, 1982; Quade and Cerling, 1995; Barry et al., 2002;
Suresh et al., 2004). 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 (Flynn et
al., 1995; Flynn, 2003; Basu, 2004). The Middle Siwaliks (Nagri, 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 Dhok Pathan type locality is characterized by the light colored sandstone with alternate clay and minor layers of conglomerates. The sediments have orange shale with less compacted gray sandstone bodies 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 upsection gradually get 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 (Behrensmeyer and Tauxe, 4
1982). 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.
Stratigraphy and Chronology
The Siwalik fauna has been regarded as one of the most important Cenozoic fossils anywhere in the world and has been studied intensely for more than a century. Several
American and Pakistani scientists did post-independence stratigraphical work in the
Siwaliks of Pakistan (Pilgrim, 1910, 1913; Colbert, 1935; Shah, 1977; 1980; Pilbeam et al., 1977; Stix, 1982; Behrensmeyer, 1987; Willis, 1993a, b; Behrensmeyer et al., 1995;
Willis and Behrensmeyer, 1995). As a result, a fine-scale magnetostratigraphic work has emerged, supplemented by vertebrate faunal zone. Because the sediments are well suited for magnetostratigraphic studies, it has been possible to establish a regional 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).
Johnson N. et al. (1982, 1985) have shown that the Potwar Siwalik sediments range between 18.3 and 0.6 Ma. During this period only one major depositional hiatus has been identified (Opdyke et al., 1979). With reference to the Geomagnetic Reversal Time Scale of Berggern et al. (1985) the ages of the Miocene Potwar formations (Kamlial, Chinji,
Nagri, and Dhok Pathan formations) and the Late Pliocene through Pleistocene formations are now fairly well understood. More recently, the geochronology of the Dhok
Pathan Formation has been refined and a reliable chronostratigraphy has been established 5 by geomagnetic polarity time scale. The magnetic polarity and stratigraphic dating has constrained the age of the Dhok Pathan Formation to between 10.1–ca. 3.5 Ma. The lower part of the Dhok Pathan Formation is dated 10.1-9.0 Ma and the upper part is considered to be dated at ca. 9.0-5.5 Ma (Cande and Kent, 1995; Barry et al., 2002). The
Dhok Pathan Rest House section yielded an age of 8–6.5 Ma. Nevertheless, paleomagnetic and biostratigraphic correlations have yielded a precise geochronology of the latest Miocene/earliest Pliocene of the Dhok Pathan (Fig. 1A) (Pickford, 1988; Barry et al., 1982, 2002).
Biostratigraphy
The Dhok Pathan Formation faunas are mainly very similar to those of the underlying sediments; thus, the suids, the carnivores and the bovids continue virtually unchanged.
Miotragocerus and Tragoportax are represented by many species; Reduncini make their first appearance together with a species very similar to Prostrepsiceros vinayaki from which Thomas (1984) accepts the age of the late Miocene. The earliest reduncine (Kobus porrecticornis) records in Eurasia and in the Siwalik Formation of India and Pakistan also date to the late Miocene (Vrba, 1995). Miotragocerus and Tragoportax appeared in the Siwaliks during the latest middle Miocene and diversified during the late Miocene, before going extinct at the end of the Miocene.
Selenoportax record indicates that it was present in the Siwaliks from the Late Middle
Miocene to the Pliocene (Akhtar, 1992; Barry et al., 2002; Khan et al., 2009a).
Nevertheless, Barry et al. (2002) recognized a maximum range of 10.3 to 7.9 Ma for this taxon. Qiu and Qiu (1995) list Selenoportax sp. from the Lufeng fauna (Chinese 6
Miocene), age of 11.1–8.0 Ma (Steininger, 1999; Flynn and Qi, 1982). Pachyportax, on the other hand, is considered as a typical Late Miocene taxon. Pachyportax occurs in the
Nagri and the Dhok Pathan zones of the Siwaliks, until Proamphibos replaces it soon after the start of the Tatrot. Hence, a Latest Miocene date around 7.0 Ma (Barry et al.,
1991) would be considered as a possible date for a fauna containing P. latidens.
Overall, the Dhok Pathan Formation faunas resemble those from the Turolian Land
Mammal ‘Age’ as defined in Europe, North Africa and West Asia. There rather restricted time range seems to fall close to the boundary between the Vallesian and the Turolian
Land Mammal ‘Ages’. So, the age of the Dhok Pathan, is considered as the late Miocene.
Barry et al. (2002) suggested that the age of the Dhok Pathan was the late Miocene, however, the cervids, the bovids, the giraffids, and the suids suggest a late Miocene to early Pliocene age. The faunal association of the locality indicates an age to date between the late Miocene and early Pliocene (Fig. 1).
Siwalik Artiodactyls
The late Miocene was a time of active speciation and origination of higher taxa in mammals (Table 1) and it is a time period that is crucial to our understanding the evolution of the extant artiodactyls (Vrba, 2000). The Dhok Pathan type locality has yielded several species of artiodactyls, perissodactyls, the oldest Siwalik cercopithecoid, carnivores, proboscideans and rodents (Table 1) clearly distinct from that found in the upper part of the Nagri Formation and its equivalents. The artiodactyls comprise almost
50% of the total faunal elements (Fig. 2).
7
Dhok Pathan Mammalian Fauna
Carnivora 20%
Rodentia Artiodactyla 3% 46% Cercopithecidae 2% Proboscidea 17% Perissodactyla 12%
Figure 2. Percentage of species represented in the Dhok Pathan mammalian fauna.
Artiodactyls are found abundantly in the late Miocene sites of the Siwaliks, Africa and
Eurasia. The fossil record of Artiodactyla from the Siwaliks (Table 1) is generally very good in comparison with that of other parts of the world and the late Miocene locality of the Dhok Pathan has yielded one of the richest fossil artiodactyl faunas from south Asia
(Lydekker, 1878; Matthew, 1929; Colbert, 1935; Pilgrim, 1926 a, b, 1937, 1939; Thomas,
1984, Pickford, 1988; Akhtar, 1992; Haile-Selassie et al., 2004; Bibi, 2007; Bibi et al.,
2009). Many Siwalik artiodactyls are represented by cranial and postcranial material and show strong relationship with Eurasian and African artiodactyls. Fossil remains have been known in the area since the nineteenth century and large mammals have been the object of several monographies (Lydekker, 1878; Matthew, 1929; Colbert, 1935; Pilgrim,
1926 a, b, 1937, 1939; Pickford, 1988).
The late Miocene artiodactyls probably descended from Lophiomerycidae in the late middle Eocene of south Asia (Guo et al., 2000). The early-middle Eocene Kuldana
Formation in northern Pakistan has yielded several forms of bunolophodont artiodactyls, all of which are endemic to the Indian subcontinent (Metais et al., 2004). The Oligocene 8
fossils from the Bugti Hills of Pakistan may also document the first appearance and
subsequent radiation of giraffes and bovids, two groups that dominate later Neogene and
modern herbivore faunas. In Pakistan true bovids may appear as early as 19.5 Ma, and the
family is certainly present at 18.5 Ma. Since they are not known earlier elsewhere, their
appearance in southern Asia may document the beginning of this very important radiation
(Barry et al., 2005). This period of time may represent a window of early representatives
of giraffids, cervids and tragulids in Pakistan. Eotragus, an early boselaphine described
from the Kamlial Formation of the Siwaliks is considered the oldest bovid of the world
(Solounias et al., 2005).
It is apparent that the artiodactyl fauna from the Dhok Pathan is dominated by bovids.
Nevertheless, there are other species of the artiodactyls in the fauna tragulids, cervids,
giraffids and suids. Out of bovids, boselaphines are abundant in the late Miocene of the
Dhok Pathan. The Siwalik was inhabited by various boselaphines during the late Miocene
and many new forms of boselaphines appeared, while their diversification was reduced in
the Early Pliocene (Khan et al., 2009a). Moya-Sola (1983) defines two generic or
suprageneric group of boselaphines comprising, the medium sized Siwalik boselaphines
genera Miotragocerus Stromer, 1928 and Tragoportax Pilgrim, 1937 and the large sized
Siwalik boselaphines genera Selenoportax and Pachyportax as well as allied forms such
as Austroportax latifrons from the middle Miocene of Europe.
Tragoportax were among the widest spread bovids in the late Miocene of the Siwaliks
(Pilgrim, 1937, 1939; Thomas, 1984; Akhtar, 1992). The genus is also widespread in
Oriental and southeastern Europe (Geraads et al., 2003; Kostopoulos, 2009). They appear
for the first time in the middle Miocene but they are best known from the Turolian 9
wherein, alongside Gazella, they are considered as the most widespread of all bovids.
This group went extinct around the end of the Miocene (Gentry, 1999). Tragoportax sp.
is present in the Turolian of the Graeco-Irano-Afganian province as well as in the
Siwaliks (Solounias, 1981; Bernor, 1986; Kohler, 1987; Gentry and Heizmann, 1996;
Kostopoulos, 2005, 2009). Tragoportax cyrenaicus at its type locality (France) is a
morphologically advanced species of the very late Miocene (Geraads, 1989: 790). Allied
species to T. cyrenaicus are found in the presumed early Pliocene locality of
Langebaanweg, south Africa (Tragoportax acrae) and in the late Miocene of the Siwaliks
(Tragoportax salmontanus; Gentry, 1999). Tragoportax salmontanus resembles closely
to the late Miocene Siwalik species Prostrepsiceros vinayaki.
Pilgrim (1937, 1939) mentioned variety of the medium sized boselaphines in the Siwaliks
and some merely based on the minor variations (e.g. Sivaceros gradiens). Thomas (1984)
referred to this group all the specimens that came from the Nagri and the Dhok Pathan
formations, and allotted the name Tragocerina. More recently, Bibi et al. (2009) assigned
the name ‘Tragoportacini’ for the Miotragocerus-Tragoportax complex.
Akhtar (1992) recovered a huge material of Tragoportacini from the type locality Dhok
Pathan and named it Tragocerus punjabiensis, Tragocerus browni, and Miotragocerus
dhokpathanensis Akhtar. According to Akhtar (1992) that Miotragocerus
dhokpathanensis is larger than Miotragocerus gluten. The species is illustrated by a single specimen opisthocranium and the characteristics of the opisthocranium allow us to compare it to Tragoportax. The skull shows all the main features of the Tragoportax (see
systematics) and it is perhaps a synonym of Tragoportax. Furthermore, many remains of 10
various taxa have been recovered from the type locality during the last four decades and
erroneously linked to Tragoportax, Tragocerus and Miotragocerus.
The revised description is proposed on the basis of the new finds (Appendix 1) and the already recovered material from the type locality Dhok Pathan. This thesis describes and interprets the entire collection of artiodactyls from the Dhok Pathan type locality
(Appendix 2), recovered during 1980 and 2009, and at present in the Palaeontology
Laboratory of the Zoology Department of the Punjab University, Lahore, Pakistan and in the Zoology Department of GC University, in Faisalabad, Pakistan.
Table 1: Mammalian faunas of Dhok Pathan. Many species are under taxaonomic revision. Artiodactyla Bovidae Tragoportax browni T. salmontanus T. perimense T. punjabicus Miotragocerus gluten Proleptobos birmanicus Selenoportax vexillarius S. lydekkeri Pachyportax latidens P. nagrii P. giganteus Eotragus sp. Gazella lydekkeri G. padriensis Taurotragus latidens Perimia falconeri Elaschistoceros khauristanensis Giraffidae Vishnutherium iravaticum Bramatherium perimense Hydaspitherium megacephalum H. grande H. magnum Giraffa punjabiensis Cervidae Rucervus simplicidens Cervus triplidens C. sivalensis C. punjabiensis C. rewati Tragulidae Dorcabune anthracotherioides D. nagrii Dorcatherium majus D. minus 11
Table 1 (Continued) Anthracotheriidae Chocromeryx silistrense Merycopotamus dissimilis
Suidae Tetraconodon magnus T. mirabilis Propotamochoerus uliginosus P. hysudricus Hippopotamodon sivalensis Hyosus punjabiensis H. tenuis Sivahyus hollandi Hippohyus lydekkeri H. grandis Sus comes S. adolescens Perissodactyla Equidae Hipparion antelopinum H. theobaldi H. perimense H. nagriensis Rhinocerotidae Aceratherium perimense A. lydekkeri Chilotherium intermedium C. blanfordi Subchilotherium intermedium Alicornops sp. Brachypotherium perimense Chalicotheriidae Chalicotherium salinum Proboscidea Dinotherium indicum D. angustidens (?) Trilophodon hasnotensis Tetralophodon falconeri T. punjabiensis Rhynchotherium chinjiensis Synconolophus dhokpathanensis S. propathanensis S. corrugatus S. ptychodus S. hasnoti Anancus perimensis Stegolophodon latidens S. cautleyi Stegodon bombifrons S. cliftii S. elephantoides
Cercopithecidae Cercopithecus hasnoti Macacus sivalensis Rodentia Rhizomys sivalensis Rhizomys sp. Hystrix sivalensis Carnivora Arctamphicyon lydekkeri Indarctos punjabiensis Promellivora punjabiensis Enhydriodon falconeri 12
Table 1 (Continued) Sivaonyx bathygnathus Vishnuictis salmontanus Ictitherium sivalense I. indicum Lycyaena macrostoma L. macrostoma-cinayaki Crocuta carnifex C. gigantean C. gigantean-latro C. mordax Mellivorodon palaeindicus, Acluropsis anneclens Paramachacrodus pilgrimi P. indicus Propontosmilus sivalensis, Felis sp.
Objectives of Present Study
The main aim of this study has been to provide the first complete documentation of the
artiodactyls found in the Dhok Pathan type locality by tackling aspects of taxonomy,
stratigraphy, palaeontology and paleoecology of the Siwaliks of Pakistan. An
ecologically important group, the artiodactyls, was selected for the study as the collected artiodactyls material presented notable diversity and thus could provide significant taxonomic, ecological, biostratigraphic and palaeoenvironmental information. Finally, the taxonomic diversity will be compiled to provide information about the palaeoenvironmental conditions that prevailed during the Late Miocene in the Dhok
Pathan
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
Punjab University, Lahore, Pakistan. The first chapter entitled “Introduction” includes mainly geography, stratigraphy, biostratigraphy and the Siwalik artiodactyls of the Dhok
Pathan. This chapter elaborates geology, stratigraphy, biostratigraphy, and chronostratigraphy of the Dhok Pathan in which a map of the Siwaliks compiled, 13 showing the correlation of the Siwalik formations and zones. The second chapter entitled
“Materials and Methods” includes methodology for the research work.
The third chapter entitled “Systematic Palaeontology” elaborates the taxonomical and morphological features of the studied material are presented. This chapter presents the taxonomical and morphological features of twenty three species of artiodactyls from the
Dhok Pathan, Middle Siwaliks, Pakistan. Consequently, repetition of description, discussion and systematics about the genera does occur in this chapter. The fourth chapter is entitled “Discussion and Conclusions” in which the Dhok Pathan artiodactyls’s palaeoecological and palaeoenvironmental requirements are interpreted. The Dhok
Pathan artiodactyls are correlated with the Eurasian and African artiodactyls in this chapter. Finally, the references and appendices are given at the end of the thesis. The thesis will be submitted for publication to journal of Vertebrate Palaeontology. One additional paper which comprised description of Selenoportax has been already published
(Appendix 3). My contribution to this paper has been the description and the discussion of the Selenoportax.
14
MATERIALS AND METHODS
The described material comprises skull, horn-cores, maxilla and mandible fragments, and isolated molars (Appendix 2). The material includes the newly collected specimens from the Dhok Pathan type locality during field trips that took place from 1980 to 2009 by the
Palaeontology team of the Punjab University, Lahore, Pakistan. In addition some of the specimens used in this study already belonged to the collections of the Palaeontology
Laboratory, University of the Punjab, Lahore, Pakistan and had been collected in the past decades.
Almost all fossil specimens were found weathering out from or in situ within the light colored sandstone with alternate clay and orange shale. Fossils were generally very well preserved. The material came from 17 localities. The specimens excavated from these localities were generally in excellent condition, with little surface damage, often complete, and sometimes in articulation. Most specimens found on erosional surfaces were also well preserved, particularly those that had not been exposed for long, as on steep, actively eroding slopes. The spatial distribution of fossil material was non-random.
Over 70% came from 6 localities of the Kundrali unit, the largest of that (localities DP11-
13) had over 65 fossils on its surface. This pattern seems typical of the Siwaliks as earlier workers already noted that fossils were typically found in pockets in the Siwaliks (Gaur,
1987; Raza et al., 2002).
Piercing instruments like chisels and geological hammers were employed for the excavation of partially embedded fossils. Careful measures were taken so as to prevent the fossils from disintegrating during excavation. Each specimen was wrapped with a cotton piece to avoid the shocks of transportation. Eventually the collected specimens 15 were brought in the laboratory for taxonomic and morphological analysis. Clay and other hardly adjoined sedimentary particles were removed with the help of fine needles and brushes. Accidentally broken fragments of the specimens were rejoined by using gums and resins such as Magic Stone, Elfy, and Fixings etc. A hand lens was used for keen observation of very small and ambiguous morphological characters.
The studied material is housed in the Paleontology Laboratory of the Zoology
Department of the Punjab University, in Lahore, Pakistan and in the Zoology Department of GC University, in Faisalabad, Pakistan. Comparisons of specimens were made with those at the Natural History Museum, London (BMNH), the Natural History Museum,
America (AMNH), the Geological Survey of Pakistan (GSP), the Geological Survey of
India (GSI), the Palaeontological collection of Government College University,
Faisalabad, Pakistan (PC-GCUF) and the specimens from the Palaeontology laboratory of the Zoology Department of the Punjab University, Lahore, Pakistan (PUPC). The specimens were catalogued by giving them a number which consists of the year of collection followed by a serial catalogue number (e.g. 83/261). Upper case letters stand for upper teeth and lower case letters for lower teeth. The selected photos of the studied taxa showing reflective morphometric features are given in the figures.
The measurements of the horn-cores are taken at the base. The anteroposterior diameter
(DAP) corresponds to the large diameter of the horn-core base and it may not be parallel to the sagittal plane. The mediolateral diameter (DT) is perpendicular to the DAP.
Measurements of the studied teeth specimens were taken with a caliper as maximum lengths, widths and height at the occlusal level and are expressed in millimeters (mm).
Tooth cusp terminology follows the nomenclature of Gentry (1994) and Pickford (1988), 16 and horn core terminology follows Pilgrim (1937, 1939), and Spassov and Geraads,
(2004).
Taxonomic Description
Careful and extensive morphometric comparison led to the taxonomical identification of twenty three artiodactyl species. The identified artiodactyl species are listed in systematic order with information on holotype, diagnosis, differential diagnosis, geographic distribution, description, comparison, and discussion.
17
SYSTEMATIC PALAEONTOLOGY
Order ARTIODACTYLA Owen, 1848
Family BOVIDAE Gray, 1821
Tribe BOSELAPHINI Knottnerus-Meyer, 1907
Genus TRAGOPORTAX Pilgrim, 1937
Type Species: Tragoportax salmontanus Pilgrim, 1937.
Generic Diagnosis (Pilgrim, 1937): Boselaphinae of moderate size; skull wide at the
frontals and occipital; face bent down on the cranial axis; occiput long or moderately
short; horn-cores rather short, with or without a slight twist, situated far apart on the
frontals, but with their inner base expanded considerably so as to form a swelling on the
frontals which sometimes bridges the entire interval between the horn-cores as an
elevated belt falling abruptly to the rear; with prominent antero-internal and postero-
external keels; keel axis very oblique to skull axis; horn-cores directed strongly backward
and moderately or strongly outward, with or without an inward curve; cross section at base subtriangular, laterally compressed, compression increasing above the base, outer face between the keels weakly convex, third angle internal, about two thirds of the way from the anterior keel to the rear, rounded; supraorbital pits small, circular; temporal ridges very strong; frontal rugose; lachrymal fossa deep; basioccipital short, subtriangular, with or without a shallow median groove; supraoccipital exposed as a rectangular area on the upper surface of the skull; upper molars quadrate, moderately 18 hypsodont, outer folds and ribs rather strong, median rib in posterior lobe weaker than in anterior lobe, entostyles present in all the molars.
Generic Diagnosis (Spassov and Geraads, 2004): Size generally large, approximately that of European Cervus elaphus. The postcornual fronto-parietal surface is a flat or slightly concave well defined depressed area, usually bordered laterally by well marked temporal ridges and caudally by a step leading to a slightly raised plateau. The basi- occipital has a longitudinal groove between the anterior and the posterior tuberosities, in the bottom of which often runs a weak sagittal crest. Adult male horn-cores are long and slender, usually curved backwards, with a triangular to subtriangular cross-section, well marked posterolateral keel and flattened lateral sides, but are less compressed than in
Miotragocerus. Anterior rugosities growing downwards from the anterior keel at the basis of horn-cores are absent or weak, and usually do not extend onto the frontal.
Demarcations (steps) on the anterior keel are often found, but are few when present.
Horn-cores have a heteronymous torsion (anti-clockwise on the right horn), so that the anterior keels first diverge in anterior view, but they re-approach towards the tips. The intercornual plateau is rather short antero-posteriorly, broad and almost rectangular between the horn-cores. The occipital is not much broader ventrally than dorsally, giving it a trapezoid (rather than triangular) outline. Teeth rather hypsodont; labial walls of upper teeth, and lingual ones of lower teeth with less accentuated ribs and styles than in
Miotragocerus. Premolars relatively shorter than in Miotragocerus. P2 short relatively to
P3, especially its anterior part, and parastyle curved backwards. P3 with lingually inflated hypocone. Metaconid of p3-p4 larger than in Miotragocerus, splayed lingually and T- shaped on p4, with an open anterior valley. 19
Stratigraphic Range: Middle Siwaliks.
Geographic Distribution: From the Vallesian/Turolian boundary (or lateVallesian?) to the end of the Turolian, perhaps earliest Pliocene in Africa. From southeastern Europe and the northern Paratethys region through Asia minor and the Middle East to Africa and the northern part of the Indian subcontinent (and possibly central Asia) (Pilgrim, 1937,
1939; Spassov and Geraads, 2004).
Tragoportax punjabicus (Pilgrim, 1910)
Type Specimen: Skull: GSI B486.
Emended Diagnosis (modified from Pilgrim 1937, 1939 and, Spassov and Geraads,
2004): A Tragoportax with moderately long, curved horn-cores with a large antero- posterior diameter, faintly twisted, slight torsion, cross-section triangular to subtriangular, well marked posterolateral and flattened sides. The postcornual fronto-parietal surface is a flat or slightly concave well defined depressed area, usually bordered laterally by well marked temporal ridges and caudally by a step leading to a slightly raised plateau. The basi-occipital has a longitudinal groove between the anterior and the posterior tuberosities, in the bottom of which often runs a weak sagittal crest. Occipital is high and basioccipital is short. Premolar series is short, upper molars hypsodont with divergent styles and prominent median ribs. P2 has a distally sloping parastyle with a short parastyle-mesostyle part and bilobed lingually. P3 has a lingually inflated hypocone. The p4 has a strong paraconid, metaconid and entoconid. The entoconid is fused with the endostylid. The anterior valley of p4 is open. 20
Studied Material: Skull: PUPC 86/149, opisthocranium. PUPC 86/150, opisthocranium.
Horn-core: PUPC 86/264, broken at the apex, left side. Upper dentition: PUPC 83/743,
isolated right P3. PUPC 87/327, isolated left P3. PUPC 00/21, isolated left P3. PUPC
00/89, right P4-M1. PUPC 83/671, isolated right M1. PUPC 87/677, isolated right M1.
PUPC 83/673, isolated left M1. PUPC 83/30, isolated left M1. PUPC 83/676, isolated
right M1. PUPC 83/674, isolated right M1. PUPC 97/13, isolated right M1. PUPC 00/80,
right M1-2. PUPC 85/242, isolated right M2. PUPC 97/06, isolated left M2. PUPC
83/603, isolated right M2. PUPC 83/10, isolated left M2. PUPC 83/666, isolated right
M2. PUPC 00/01, isolated left M2. PUPC 83/693, isolated right M2. PUPC 08/126, palatine parts of both maxillae with right P3-M3 and left P2-M3. PUPC 83/275, right maxillary fragment with P3-M3. PUPC 97/10, right maxillary fragment with M1-M3.
PUPC 83/665, isolated left M3. PUPC 97/7, isolated right M3. PUPC 83/688, isolated right M3.
Lower dentition: PUPC 83/712, isolated left p2. PUPC 97/12, left mandible fragment with p2-p4. PUPC 83/204, isolated left p3. PUPC 87/324, isolated left p3. PUPC 83/711, isolated left p4. PUPC 68/237, isolated left p4. PUPC 97/3, isolated right m1. PUPC
87/282, isolated left m1. PUPC 86/26, isolated left m1. PUPC 83/700, isolated right m1.
PUPC 86/25, isolated left m2. PUPC 83/617, isolated left m2. PUPC 99/28, isolated right m2. PUPC 83/701, isolated left m2. PUPC 83/680, isolated left m3. PUPC 83/767, isolated left m3. PUPC 83/277, left mandible fragment with m3.
21
Description
Skull
PUPC 86/149 and PUPC 86/150 opisthocraniums are incomplete specimens having only
the occipital region, right occipital condyle, paraoccipital process, parietals, and
basisphenoid (Fig. 3). PUPC 86/150 opisthocranium (Fig. 3(2)) is more fragile than
PUPC 86/149 (Fig. 3(1)). The facial part, horn-core, one occipital condyle and dentition
are missing. The sutures separating the various bones are not visible. From the occipital
foramen to the sphenoid, a continuous groove runs along the basioccipital, often with a weak sagittal crest in its middle. The groove is very prominent in PUPC 86/150 and the sagittal crest is getting weak towards the posterior tuberosities (Fig. 3(2)). The anterior tuberosities are fairly wide apart and not very large while the posterior tuberosities are large and expanded laterally. The parietal surface is a flat and slightly concave well defined depressed area, bordered laterally by well marked temporal ridges and caudally by a step leading to a slightly raised plateau. The temporal outer ridge runs back less obliquely and finally bends round by the side of the supraoccipital to join the lambdoid crest. The basioccipital is subtriangular in shape and the paraoccipital process points ventrally. The comparative measurements of the opisthocraniums are provided in table 2.
Horn-core
The horn-core is fragment and incomplete, lost upper part but the basal part is complete
(Fig. 4). The horn-core is moderately long, compressed laterally. It is curved backward, weak torsion and faintly twisted. The lateral face of the horn-core is flat while the internal one is slightly convex. It has a prominent antero-internal keel and a rounded postero- external keel. The cross section of the horn-core at the base is triangular. The medio- 22
lateral compression increases from the base to the top. Weak discontinuous longitudinal
grooves run along the horn-core surface. The total length measured along the anterior
face is 143 mm and along the posterior face is 140 mm. The transverse diameter at the base is 31 mm and antero-posterior diameter at the base is 53 mm. The mean index DT ×
100/DAP is 58.4 at the base.
Upper dentition
The premolar row is relatively shorter than the molar row. PUPC 08/126 is only one sample which shows complete cheek teeth series (Fig. 5(4)). The length of the premolar series is 32.5 mm and the length of the molar series is 52 mm in PUPC 08/126. P2 is
similar to P3 but it is short to P3. P2 has a distally sloping parastyle with a short
parastyle-mesostyle part and bilobed lingually. It has a weak hypocone. P3 is larger than
P2. It has a narrow but very pronounced parastyle. P3 has a broad and strong posterior
fold with a mesostyle anteriorly. P3 has a lingually inflated hypocone. P4 is a horse shoe shaped with strong anterior and posterior folds. A slight median rib is present. A very prominent and divergent parastyle is present in P4. A metastyle is moderately present. A short spur projects in the central fossette of the posterior side of the inner crescent in P4.
The occlusal outline of the premolar is nearly rectangular, being much broader transversely than long. A sharp praeprotocrista unites a protocone with a parastyle.
The molars are quadrate and hypsodont. The enamel is finely rugose. The molar row is longer than the premolar row. The small entostyle is present in all molars lingually. The entostyles are slightly shifted towards the hypocone. The parastyle and mesostyle are stronger than the metastyle. The metastyle is strong and posteriorly extended on M3. The anterior median rib is more prominent than the posterior one. A faint cingulum is present 23
on the antero-internal and postero-internal surface of the molars. The main morphological
features of the upper molars are the well projected mesostyle, the wider hypocone to the
protocone comparatively and strong parastyle directed anteriorly (Fig. 5(4-5)).
Lower dentition
The lower premolars and molars are rugose (Fig. 5(6-7)). The paraconid of p2 is weaker than p3. The metaconid of p3 is backwardly directed. The entoconid is stronger than the metaconid. The paraconid of p3 is stronger than parastylid and placed antero-posterior axis of the premolar. The p4 is extended antero-posteriorly and similar to p3 (Fig. 5(6)).
The p4 has a strong paraconid, metaconid and entoconid. The entoconid is fused with the endostylid. The anterior valley of p4 is open. A vestigial ectostylid is present in the molars. The hypoconulid of m3 is rather large and variable. The comparative measurements of teeth are provided in table 3.
24
Table 2: Comparison of the cranial measurements (mm) of the Siwalik Tragoportax. Description T. T. T. browni T. browni T. browni T. perimensis T. cf. punjabicus punjabicus (PUPC (Pilgrim, (Akhtar, (Pilgrim, salmontanus (PUPC (Pilgrim, 86/204, in 1939) 1992) 1939) (PUPC 86/149, in 1939) this thesis) 93/20, in this this thesis) thesis) Breadth of skull at orbits - 109 160 117 - ?125 - Breadth of skull at - 93 180 87 86 - - mastoid Height of occipital - - 100 - - - - Width of the braincase 82 70 111 70 81 ?71 - Distance between outer - - 96 - - - - edges of the occipital condyles Antero-posterior - 71 65 65 - 58 33 diameter of the horn core at base Transverse diameter of - 36 42 36 - 37 25.4 the horn core at base Distance between fronto- - 81 98 - - - - parietal suture to summit of occipital crest Distance between parieto- - 24 30 - - - - supraoccipital suture to summit of occipital crest
25
Table 3: Comparative measurements of the cheek teeth of T. punjabicus in mm (millimeters). * The studied specimens. Referred data are taken from Pilgrim (1939) and Akhtar (1992). Number Nature/Position Length Width W/L ratio
PUPC 83/743* rP3 15.0 13.0 0.86 PUPC 87/327* lP3 13.0 12.0 0.92 PUPC 00/21* lP3 14.0 12.0 0.85 PUPC 00/89* rP4 12.0 13.0 1.08 rM1 15.0 16.0 1.06 PUPC 83/671* rM1 17.4 16.2 0.93 PUPC 87/677* rM1 18.0 16.0 0.88 PUPC 83/673* lM1 18.0 18.0 1.00 PUPC 83/30* lM1 15.0 16.0 1.06 PUPC 83/676* rM1 16.4 15.0 0.91 PUPC 83/674* rM1 16.4 16.4 1.00 PUPC 97/13* rM1 17.0 14.0 0.82 PUPC 00/80* rM1 17.0 16.0 0.94 PUPC 85/242* rM2 18.0 16.5 0.91 PUPC 97/06* lM2 18.0 17.0 0.94 PUPC 83/603* rM2 19.0 16.0 0.84 PUPC 83/10* lM2 19.0 15.3 0.80 PUPC 83/666* rM2 18.8 17.5 0.93 PUPC 00/01* lM2 17.0 17.0 1.00 PUPC 83/693* rM2 16.8 16.0 0.95 PUPC 08/126* lP2 8.00 6.00 0.75 lP3 13.0 9.40 0.72 lP4 10.5 12.0 1.14 lM1 16.3 14.4 0.88 lM2 18.0 15.0 0.83 lM3 18.3 13.5 0.73 PUPC 83/275* rP3 14.0 13.0 0.92 rP4 11.0 13.7 1.2 rM1 17.0 16.0 0.94 rM2 18.0 17.0 0.94 rM3 18.4 16.5 0.89 PUPC 97/10* rM1 15.0 15.5 1.03 rM2 18.0 16.0 0.88 rM3 21.0 16.0 0.76 PUPC 83/665* lM3 18.0 17.0 0.94 PUPC 97/7* rM3 18.5 17.0 0.91 PUPC 83/688* rM3 19.0 15.0 0.78 PUPC 83/712* lp2 12.0 8.00 0.66 PUPC 97/12* lp2 11.0 5.00 0.45 26
Table 3 (Continued) lp3 14.0 7.00 0.50 lp4 15.0 10.0 0.66 PUPC 83/204* lp3 13.0 7.40 0.56 PUPC 87/324* lp3 14.4 3.40 0.23 PUPC 83/711* lp4 15.0 8.00 0.53 PUPC 68/237* lp4 15.0 8.00 0.53 PUPC 97/03* rm1 15.0 10.0 0.66 PUPC 87/282* lm1 17.0 10.0 0.58 PUPC 86/26* lm1 15.5 11.0 0.70 PUPC 83/700* rm1 15.0 9.00 0.60 PUPC 86/25* lm2 18.5 11.0 0.59 PUPC 83/617* lm2 18.0 10.5 0.58 PUPC 99/28* rm2 19.0 11.0 0.57 PUPC 83/701* lm2 17.0 12.0 0.70 PUPC 83/680* lm3 25.0 11.0 0.44 PUPC 83/767* lm3 22.0 10.0 0.45 PUPC 83/277* lm3 24.0 11.0 0.45 GSI B486 P2 17.0 11.0 0.64 P3 14.5 12.0 0.82 P4 11.0 15.5 1.40 M1 18.0 18.0 1.00 M2 20.0 20.0 1.00 M3 20.0 18.5 0.92 GSI B574 M1 18.0 19.0 1.05 M2 20.5 22.0 1.07 M3 21.0 20.5 0.97 GSI B563 p2 12.5 7.00 0.56 m1 19.0 12.5 0.65 m2 21.0 13.0 0.61 m3 30.0 13.5 0.45 GSI B564 p2 13.0 7.00 0.53 p3 16.0 8.50 0.53 p4 17.5 10.0 0.57 m1 17.5 12.0 0.68 m2 20.5 14.0 0.68 m3 26.0 13.5 0.51
27
Comparison
The general morphological characters of the available opisthocraniums, horn-core and dentitions show clear affinities with Tragoportax (Spassov and Geraads, 2004; Bibi and
Gulec, 2008; Kostopoulos, 2005). The material (Fig. 3-5) is very close in morphology as
well as in dimensions to the material from the already described material of the type
locality (Pilgrim, 1937, 1939). The skull (Fig. 3) shows the Tragoportax postcornual and basioccipital features and basal horn-core cross section close to Tragoportax (Spassov and Geraads, 2004; page 349). The skulls are almost identical to the type specimens of T. punjabicus and sharing in common a similar flat postcornual fronto-parietal surface with slightly concave well defined depressed area, well marked temporal ridges and caudally by a step leading to a slightly raised plateau and basi-occipital has a longitudinal groove between the anterior and the posterior tuberosities, in the bottom of which often runs a weak sagittal crest. The basal horn-core cross section is close to T. punjabicus. The dental proportions of the T. punjabicus fall within the range of the type specimens (Fig. 10), being clearly distinct from those of large Siwalik Tragoportax, T. browni and small
Tragoportax, T. salmontanus (Table 3-4). It is therefore no doubt for the inclusion of the studied material in T. punjabicus (Pilgrim, 1937, 1939). The species is also present in the
Miocene of Eurasia (Kostopoulos, 2009; Bibi and Gulec, 2008). It differs from T. browni by the small size of skull, horn-cores and dentitions (e.g. see Fig. 10). 28
Figure 3. Tragoportax punjabicus. 1. PUPC 86/149 opisthocranium: a) dorsal view, b) ventral view, c) occipital view. 2. PUPC 86/150 opisthocranium: ventral view. Scale bar equals 30 mm total. 29
Figure 4. Tragoportax punjabicus. 3. PUPC 86/264 left horn core: a) lateral view, b) medial view. Scale bar equals 30 mm total.
30
Figure 5. Tragoportax punjabicus. 4. PUPC 08/126 palatine parts of both maxillae with right P3-M3 and left P2-M3: occlusal view. 5. PUPC 83/275 right maxillary fragment with P3-M3. 6. PUPC 97/12 left mandible fragment with p2-p4. 7. PUPC 83/277 left mandible fragment with m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. 31
Tragoportax browni (Pilgrim, 1937)
Type Specimen: Skull: AMNH 19662.
Emended Diagnosis (modified from Pilgrim, 1937, 1939): A Tragoportax of large size.
The size is larger than Eurasian T. curvicornis and the Siwalik T. punjabicus. Skull is broader at the orbits than in many other species. Horn-cores are long but with a smaller antero-posterior diameter than other Tragoportax species, tilted backwards and strongly curved. Dentitions are hypsodont and larger than T. punjabicus.
Studied Material: Opisthocranium: PUPC 86/204. Horn-cores: PC-GCUF 09/11, almost
complete, side indeterminate. PUPC 87/1, fragment, side indeterminate. Upper dentition:
PUPC 87/281, isolated left P3. PC-GCUF 09/42, isolated right P3. PC-GCUF 09/43, isolated right P4. PUPC 83/689, isolated left M1. PUPC 86/151, left maxillary fragment with M1-M2. PUPC 86/29, left maxillary fragment with M1-M2. PUPC 83/861, left maxillary fragment with M1-M2. PUPC 97/80, isolated left M2. PUPC 83/662, isolated right M2. PUPC 87/324, right maxillary fragment with M2-M3. PUPC 83/283, right maxillary fragment with M2-M3. Lower dentition: PUPC 99/27, right mandible fragment with m1-m2. PUPC 83/265, left mandible fragment with m2-m3. PUPC 83/269, right
mandible fragment with m2-m3. PUPC 97/11, left mandible fragment with m3.
Description
Skull
PUPC 86/204 opisthocranium is a fragment and incomplete specimen retaining only the
occipital region, one occipital condyle, paraoccipital process, parietals, frontals, base of
the right horn-core and basisphenoid (Fig. 6(1)). The opisthocranium is not compressed 32
dorso-ventrally but the facial part, horn-core, one occipital condyle and dentition are
missing. The sutures separating the various bones are pretty visible indicating relatively
young animal. The suture between the parietal and the supraoccipital is more prominent
than the parietal bones and the coronal suture. The intercornual area is rectangular and
short antero-posteriorly. The fronto-parietal postcornual area is depressed and slightly
concaves (Fig. 6(1a)). The occipital surface is rectangular, its dorsal part being broad.
From the occipital foramen to the sphenoid, a continuous groove runs along the
basioccipital, often with a weak sagittal crest in its middle. The opisthocranium is wide at
the orbits and the occipital.
The horn-cores are situated above the orbits and the cross section of the horn-core at the
base is subtriangular. There is a shallow depression on the frontals behind each horn-
core. The temporal ridges are strongly developed. They bifurcate a little behind the
fronto-parietal suture. The inner ridge runs straight backwards and joins its fellow in the
middle of the suture between the parietal and supraoccipital, enclosing a flat elevated
area. The outer ridge runs back less obliquely and finally bends round by the side of the
supraoccipital to join the lambdoid crest.
The supraoccipital occupies a wide area of the skull dorsally i.e. 30 mm. The occipital is
slightly concave in vertical direction above the foramen magnum. Nevertheless, the knob
which forms the top of the occipital crest is prominent and wide and the occipital condyle
projects to the rear of the occipital crest (Table 3). The foramen magnum is large and the
lamboidal crest is strongly developed. The basioccipital is subtriangular in shape. The anterior tuberosities are fairly wide apart and not very large while the posterior 33
tuberosities are large and expanded laterally. A faint median crest is present between the
tuberosities. The auditory bulla is long, laterally compressed and considerably inflated.
Horn-core
The horn-core is long, curved, inclined backward and almost complete. The horn-core
divergence decreases towards the tip. The torsion is weak and the basal cross section is
subtriangular. The horn-core is compressed laterally and straight in its lower portion (Fig.
7(2)). The antero-posterior diameter is large with a sharp anterior keel and a base with
rounded corners. The antero-internal and postero-external keels are prominent. The
surface is regularly ornamented by numerous longitudinal furrows that are sharper at the
posterior than on the anterior. The longitudinal furrows indicate torsion. The concavity at
antero-internal is prominent towards the apex and the tip of the horn-core is obtuse. The
total length of PC-GCUF 09/11 measured along the anterior face is 355 mm. The
transverse diameter at the base is 32 mm and antero-posterior diameter at the base is 57
mm. The mean index DT × 100/DAP is 56 at the base.
Dentition
The P3 has inflated hypocone. The paracone and the parastyle are strong. It has a narrow
and prominent anterior fold, a broad and strong posterior fold. The median rib is placed
anteriorly. The P4 is horse shoe-shaped with a rounded lingual wall. The molars are
moderately high crowned. The upper molars are rugose with shining enamel. A
rudimentary entostyle is present on the M1-M3 (Fig. 8(3)). A weak cingulum is present in
the upper molars. The styles are very strong. The anterior median rib is more prominent
than the posterior one. A spur is present in the posterior cavity. The lower molars have
wrinkled enamel. The rugosity is very prominent in the lower molars. An ectostylid 34
appears on the m1-m3 and goes up to the centre of the transverse valley (Fig. 8(4)). The
hypoconulid is rather large and slight convex lingually, have tendency to be elongated
(Table 4).
Table 4: Comparative measurements of the cheek teeth of T. browni in mm (millimeters). * The studied specimens. Referred data are taken from Pilgrim (1937) and Akhtar (1992). Number Nature/Position Length Width W/L ratio PUPC 87/281* lP3 13.0 11.0 0.84 PC-GCUF 09/42* rP3 13.0 11.0 0.84 PC-GCUF 09/43* rP4 12.0 13.0 1.08 PUPC 83/689* lM1 18.0 18.0 1.00 PUPC 86/151* lM1 19.0 18.0 0.94 lM2 21.0 17.5 0.83 PUPC 86/29* lM1 18.0 16.0 0.88 lM2 20.5 18.0 0.87 PUPC 83/861* lM1 16.0 16.0 1.00 lM2 20.0 16.0 0.80 PUPC 97/80* lM2 21.0 19.0 0.90 PUPC 83/662* rM2 20.0 18.0 0.90 PUPC 87/324* rM2 18.0 15.0 0.83 rM3 20.0 16.0 0.80 PUPC 83/283* rM2 20.0 15.3 0.76 rM3 20.4 20.0 0.98 PUPC 99/27* rm1 16.0 12.0 0.75 rm2 18.0 13.0 0.72 PUPC 83/265* lm2 17.0 10.4 0.61 lm3 22.0 11.0 0.50 PUPC 83/269* rm2 17.0 12.0 0.70 rm3 23.0 11.0 0.47 PUPC 97/11* lm3 26.0 11.0 0.42 AMNH 19662 P2 15.5 12.5 0.80 P3 15.0 17.0 1.13 P4 12.5 17.0 1.36 M1 18.0 18.0 1.00 M2 20.0 20.0 1.00 M3 21.0 20.0 0.95 AMNH 29884 p2 11.5 6.00 0.52 p3 15.0 8.00 0.53 p4 15.5 9.00 0.58 m1 16.0 11.0 0.68 m2 19.0 12.0 0.63 m3 25.5 12.0 0.47 Comparison 35
The referred material includes skull, horn-cores and dentitions. The dentition comprises
both mandibles and maxillae. The specimens represent the typical characteristics of
Tragoportax. The studied skull postcornual and basioccipital features and the basal horn-
core cross section close to Tragoportax (Spassov and Geraads, 2004). The skull shows the Tragoportax postcornual and basioccipital features and basal horn-core cross section close to Tragoportax. The skull has large dimensions and shows all the main features of
Tragoportax. The specimens are distinguished from T. punjabicus by their greater size
(Table 2-4) and they are pretty match of T. browni, another large Siwalik Tragoportax species. Tragoportax browni has no distal demarcation on the horn-cores. The horn-core is large one to size of T. cyrenaicus of Abu Dhabi (Gentry, 1999) but it is less divergent and has no distal demarcation. The dental and horn-core proportions of the T. browni fall within the range of the type specimens, being clearly distinct from those of other Siwalik
Tragoportax, T. punjabicus and T. salmontanus (Fig. 10). It is therefore, the studied materials close to T. browni and exhibit large Siwalik specimens of Tragoportax. It differs from T. punjabicus and T. salmontanus by the large size of skull, horn-cores and dentitions (e.g. see Fig. 10).
36
Figure 6. Tragoportax browni. 1. PUPC 86/204 opisthocranium: a) dorsal view, b) ventral view, c) occipital view. Scale bar equals 30 mm total. 37
Figure 7. Tragoportax browni. 2. PC-GCUF 09/11 horn core, side indeterminate: a) anterior view, b) posterior view, c) medial view, d) lateral view. Scale bar equals 30 mm
total.
38
Figure 8. Tragoportax browni. 3. PUPC 86/151 left maxillary fragment with M1-M2. 4.
PUPC 83/265 left mandible fragment with m2-m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
39
Tragoportax cf. salmontanus Pilgrim, 1937
Type Specimen: Skull: AMNH 19467.
Diagnosis (Pilgrim, 1937): A Tragoportax of small size; with short, slightly twisted horn-cores, having a slight inward curve, slightly divergent, the angle of divergence being about 40°; keels of the horn-cores meeting, if prolonged, at a right angle between the supraorbital foramina; horn-cores much expanded internally at the base and with a strong frontal swelling between them; occiput short and relatively broad, the length of the temporal fossa being to the width of the brain case in the ratio of about 0.93 to 1.
Studied Material: Horn-core: PUPC 93/20, incomplete, left side.
Description
The horn-core is relatively small, almost complete at the base and it is broken at the centre (Fig. 9). The anterior keel appears at the base. The lateral face of the horn-core is flat and the internal one slightly convex. The anterior and the posterior surfaces of the horn-core have longitudinal furrows. The horn-core has a very slight clockwise twist. The cross section is equilateral triangle at the base and look to become compressed laterally towards the apex. The transverse diameter of PUPC 93/20 at the base is 25.4 mm and antero-posterior diameter at the base is 33 mm giving its mean index DT × 100/DAP
77%.
Comparison
The cross section of the horn-core is typically of Tragoportax in having triangular shape and prominent anterior keel. The horn-core is shorter than in T. punjabicus and T. browni 40
(e.g. see Table 2 and Fig. 10). Tragoportax salmontanus has short horn-cores and the antero-posterior diameter is less (Pilgrim, 1937). The horn-core shape, size and the cross section is almost identical with T. salamontanus (Pilgrim, 1937, 1939; Kostopolous,
2009). However, the material is imperfect, based only on the horn-core and more perfect material will be required for the confirmation. Therefore, T. cf. salamontanus can be reasonably assigned for the horn-core, based on the morphometric characteristic. The horn-core of European Hadjidimovo Tragoportax are longer, less inclined and not so twisted than those of T. salmontanus from the Siwaliks (Pilgrim, 1939).
Figure 9. Tragoportax cf. salmontanus. 1. PUPC 93/20 left horn core: a) anterior view, b) posterior view, c) medial view. Scale bar equals 30 mm total.
41
Discussion
Boselaphines become especially diversified in the Siwaliks. The genus Tragoportax created by Pilgrim (1937) for the Siwalik species T. salmontanus. The sample of
Tragoportax exhibits fairly large range of size and morphology (Fig. 10). The two large
Siwalik species T. punjabicus and T. browni shows overlapping in size (Fig. 10).
However, the diagnostic features are enough to distinguish them for each other.
Tragoportax browni might be synonym of T. punjabicus as proposed by Kostopoulos
(2009) but it needs a more perfect material to confirm this hypothesis. Tragoportax is widespread in much of Europe and southern Asia during the Turolian (Bibi and Gulec,
2008) and recently, named the group Tragoportacini (Bibi et al., 2009). The different species of Tragoportax are best distinguished on the basis of horn-core characters. The horn-cores of the poorly known T. perimensis from the Middle Siwaliks (Pilgrim 1939) are much shorter. Another poorly known species from the Siwaliks is T. islami.
Tragoportax islami and T. perimensis are a possible synonym of T. salmontanus. The difference in horn-core curvature and in size is probably due to individual variability
(Kostopoulos, 2009).
Three species of Tragoportax could be identical from the Siwaliks, based on the present investigation, one is small T. salmontanus and the second is intermediate T. punjabicus and the third is large T. browni. There is variability in diagnostic characters of
Tragoportax and we can include already described skull Miotragocerus dhokpathanensis
Akhtar, 1992 in T. browni. These species could be identical. The differences mentioned by Akhtar (1992) are metrical. Akhtar (1992) was not considered variability of
Tragoportax. The skull is very close in morphology as well as in dimensions to the 42 already described material of Tragoportax (Table 2). Therefore, Miotragocerus dhokpathanensis is a possible synonym of T. browni. The skull is identical with the skull of T. browni and represents the same species.
T. punjabicus T. browni T. perimensis T. cf. salmontanus
Horn-core proportions of the Siwalik Tragoportax P2 14 12 80 10 60 8 6 40 Width DAP 4 20 2 0 0 0 2 4 6 8 10 12 14 16 18 0 1020304050 Length DT P4 P3 18 20 16 14 15 12 10 10 8 Width Width 6 5 4 2 0 0 0 2 4 6 8 10121416 0246810121416 Length Length
M1 M2 20 24 18 22 16 20 18 14 16 12 14 10 12
Width 8 Width 10 6 8 6 4 4 2 2 0 0 0 2 4 6 8 101214161820 0 2 4 6 8 10 12 14 16 18 20 22 24 Length Length
M3 p2 22 9 20 8 18 7 16 6 14 12 5 10 Width 4 8 Width 3 6 4 2 2 1 0 0 0 2 4 6 8 10 12 14 16 18 20 22 24 0246810121416 Length Length 43
T. punjabicus T. browni T. perimensis T. cf. salmontanus
p3 p4 10 12 8 10
8 6 6 Width 4 Widt h 4 2 2 0 0 0 2 4 6 8 10 12 14 16 0246810121416 Length Length
m1 m2 18 18 16 16 14 14 12 12 10 10 8
8 Width Width 6 6 4 4 2 2 0 0 0 2 4 6 8 10 12 14 16 18 20 0 2 4 6 8 10 12 14 16 18 20 Length Length
m3 18 16 14 12 10 8 Width 6 4 2 0 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 Length
Figure 10. Scatter diagram showing horn-cores and dental proportions of the Dhok Pathan Tragoportax. Referred data are taken from Pilgrim (1937, 1939) and Akhtar (1992).
44
Genus MIOTRAGOCERUS Stromer, 1928
Type Species: Miotragocerus monacensis Stromer, 1928.
Generic Diagnosis (Solounias, 1981): Horn-cores triangular in cross section; above the
orbits; not particularly compressed; converging anteriorly and forming a pronounced
frontal buttress; not particularly twisted but with twist restricted to tips; gently diverging
with tips neither turned in nor out; posterior grooves; anterior keel blunt, stopping about
two-thirds from base forming at least one characteristic anterior demarcation (bump);
horn-core thinner and rounder in cross section from point at which keel stops to tip; horn-
core axis more vertical and bases broader anteroposteriorly, pedicals more poorly formed than in Tragoportax. Anterior keel blunter, often with several distinct growth bumps in males; less blunt, with a single bump in females. No postcornual pits; frontals strongly
depressed behind horns; basicranium not particularly angled in relation to palate;
preorbital fossa deep; supraorbital pits small, variable in number and position. Premolars
longer in relation to the molars than in Tragoportax; P2 long; P3 with small hypocone in
relation to protocone; upper molar central cavities connect at mid-wear; entostyle small
on upper molars. p4 cavity between paraconid and metaconid open; p4 paraconid tends to
be larger than parastylid. Differs from Mesembriportax in having less sinused frontals;
differs from Protragocerus in having longer, more compressed horn-cores with an
anterior keel.
Generic Diagnosis (Spassov and Geraads, 2004): (The shape of horn-cores and those
of associated structures are those of adult males). Size small (about that of fallow deer).
The postcornual area of the skull is not depressed or raised as a low plateau.
Basioccipital, definitely known in the subgenus Pikermicerus, without median 45
longitudinal groove between the anterior and the posterior tuberosities, but with a faint
sagittal keel. Strong temporal crests (at least in males) in early forms (Miotragocerus),
weaker in more recent ones (Pikermicerus). Horn-cores moderately long to long in early
forms and short in later ones, mediolaterally compressed, with flattened lateral and
medial surfaces. Sharp anterior keel, but postero-lateral keel absent or poorly marked,
and posterior face not well delimited. The section is therefore sub-elliptic. Anterior
rugosities at base of horn-cores usually strong, extending onto the frontal along the keel,
which often has several demarcations (steps) along its course. In front view, due to a
slight torsion of the horn-cores bases, the keels are often slightly convergent upwards in
the basal portion, then diverge towards the tips. The intercornual area is much longer than
broad, especially narrow anteriorly. The occipital surface is high, much broader basally
than at its top. Teeth brachydont, with strongly folded walls. Compared to Tragoportax,
well documented forms have a long premolar row, with especially long P2 compared to
P3, due to lengthening of its anterior part. Hypocone of P3 poorly expanded lingually.
Metaconid of p3-p4 weak, anterior valley with incipient lingual wall.
Stratigraphic Range: Lower and Middle Siwaliks.
Geographic Distribution: Europe and south Asia (Spassov and Geraads, 2004).
Miotragocerus large sp.
Abbreviated Diagnosis (Pilgrim, 1939; Solounias, 1981; Spassov and Geraads,
2004): The premolars are long in relation to the molars. The premolars are as long as in
Tragoportax which is a relatively larger animal. P2 has a well developed anterior metastyle unlike Tragoportax. The protocone of P3 is medially situated unlike 46
Tragoportax. The upper molar central cavities connect at mid-wear and the entostyles are smaller than in Tragoportax. The lower dentition is more primitive than Tragoportax.
The p4 cavity between the paraconid and the metaconid is open and therefore p4 is similar to p3. The lower molars have transversely situated protoconids and hypoconids and the hypoconulid of m3 is large.
Studied Material: Upper dentition: PUPC 83/275A, isolated right P2. PUPC 86/33, isolated right P3. PUPC 86/39, isolated left P3. PUPC 09/44, right maxillary ramus with
P2-P4. PUPC 83/209, left maxillary ramus with P2-M1. PUPC 87/02 and PUPC 86/308, right maxillary ramus with DP4-M1. PUPC 87/246, left maxillary ramus with DP3-M2.
PUPC 87/115, left maxillary ramus with DP4-M2. PUPC 87/116 and PUPC 87/117, right and left maxillary rami with M1-M2. Lower dentition: PUPC 88/740, left mandibular ramus with p2. PUPC 83/703, isolated left p2. PUPC 83/708 and PUPC 83/709, isolated right and left p3s. PUPC 83/742 and PUPC 87/294, isolated right and left p4s. PUPC
93/277, left mandibular ramus with p4-m1. PUPC 09/86, left mandibular ramus with p4- m3. PUPC 96/05, left mandibular ramus with p4-m1. PUPC 83/277, right mandibular ramus with m2 and broken m3.
Description
Upper dentition
The deciduous molars are trilobed with thin enamel. The styles are bulky and divergent comparatively (Fig. 11(2)). The antero-posterior length of the deciduous molar is considerably greater than the transverse diameter (Table 5). Height of the lobes increases antero-posteriorly, anterior one being the smallest. The enamel is thin and rugose. The 47
ribs are strongly broad and highly projected. The rib of the third lobe is slightly tilted
forward near the apex. A low and broad tubercle is present between the first and the
second lobes on the lingual side. The fossette of the first lobe is simpler and wider than
those of the second and the third lobes. The labial side of the third lobe is broad at the
base, becoming abruptly narrow and pointed at the apex.
The P2 is longer than the P3 and the P4 (Fig. 11(1)). The P2 is bilobed with strongly
developed paracone-parastyle-protocone complex. The paracone rib is strong and placed
anteriorly. The P2 is more moralized than the P3. The P3 is quadrangular and less
molarized than P2 (Fig. 11(1)). The hypocone of P3 is poorly expended lingually. A
weak furrow is present lingually, separating the hypocone from the protocone. The
protruding paracone rib and the parastyle develop a narrow furrow anteriorly. The P4 has
a rounded lingual wall and centrally placed median rib. The P4 has a weaker styles and a weaker paracone than P3.
The molars are brachydont. The labial styles and ribs are well developed and strongly
projected in upper molars (Fig. 11(2)). The anterior median rib is well projected than the
posterior one. The posterior one is slightly flat. The mesostyle is robust pillar like
structure. The styles are divergent. A small tubercle like entostyle is present in upper
molars with strong folded walls.
Lower dentition
The lower dentition is brachydont with shining enamel (Fig. 11(3)). The lower premolars
show tendency of molarization, lingually. The p2 is simple with a weak paraconid and
strong medio-lingual cuspid. The weak praeprotoconulidcristid is present which is hardly 48
distinguished from the fine postprotoconulidcristid. The postmetacristid is elongated and
directs strongly backwards, tending to reach the postentocristid.
The p3 has stronger paraconid than parastylid, situated vertically antero-posterior axis of
the tooth but projected posteriorly towards the base. The preprotoconulidcristid
distinguishes from the postprotoconulidcristid. The triangular metaconid is placed just
behind the protoconid. It has flat lingual wall and enlarges from the tip to the base,
tending to close the medial valley. The postentocristid and the posthypocristid are
elongated and fused together lingually, forming a narrow, shallow and closed lingually posterior valley. The anterior valley is narrow while the medial one is wide. The paraconid is separated from the parastylid. The metaconid of p3 is free, elongated and backwardly directed. The entoconid is subtriangular and stronger than hypoconid. The hypoconid is separated from the protoconid through a labial groove.
The p4 is similar to p3 but it is larger than p3. The paraconid of p4 is strong and the
metaconid is extended anteroposteriorly. The entoconid of p4 is strong, fused rapidly
with the endostylid. The hypoconid and the protoconid are angular labially. The labial
groove between the protoconid and the hypoconid is moderately deep. Both the prae- and post-metacristid are more developed and extend anteroposteriorly, tending to close the medial valley. The lower molars have a weak anterior transverse flange and a small ectostylid that increases from m1 to m3. The talonid of m3 is single-tubercled. The metaconid and entoconid are equally developed. The protoconid and the hypoconid are slightly constricted labially and the metastylid is stronger than the parastylid and the entostylid (Table 5).
49
Table 5: Comparative measurements of the cheek teeth of Miotragocerus large sp. and Miotragocerus gluten (Pilgrim, 1937) in mm (millimeters). * The studied specimens. Referred data are taken from Pilgrim (1937). Taxa Number Nature/Position Length Width W/L ratio M. large sp. PUPC 83/275A* rP2 16.0 10.0 0.62 PUPC 86/33* rP3 14.3 10.6 0.74 PUPC 86/39* lP3 13.0 11.5 0.88 PUPC 09/44* rP2 15.5 - - rP3 15.0 12.0 0.80 rP4 13.0 15.0 1.15 PUPC 83/209* lP2 17.0 11.0 0.64 lP3 14.0 11.0 0.78 lP4 11.0 13.0 1.18 lM1 13.0 14.0 1.07 PUPC 87/02* rDP4 19.0 13.0 0.68 rM1 18.0 16.0 0.88 PUPC 86/308* rDP4 20.0 13.4 0.67 rM1 19.8 18.0 0.90 PUPC 87/246* lDP3 17.0 9.40 0.55 lDP4 19.0 14.0 0.73 lM1 19.4 17.0 0.87 lM2 22.4 17.0 0.75 PUPC 87/115* lDP4 18.0 13.0 0.72 lM1 19.0 15.5 0.81 lM2 20.5 17.0 0.82 PUPC 87/116* lM1 20.0 19.0 0.95 lM2 21.0 19.6 0.93 PUPC 88/740* lp2 12.6 6.50 0.51 PUPC 83/703* lp2 13.6 7.00 0.51 PUPC 83/708* rp3 15.2 6.30 0.41 PUPC 83/709* lp3 14.0 8.00 0.57 PUPC 83/742* rp4 17.0 8.40 0.49 PUPC 87/294* lp4 16.0 8.70 0.54 PUPC 93/277* lp4 15.0 8.00 0.53 50
Table 5 (Continued) lm1 14.0 10.3 0.73 PUPC 09/86* lp4 14.7 9.00 0.61 lm1 16.0 11.5 0.71 lm2 17.0 11.0 0.64 lm3 25.4 11.4 0.44 PUPC 96/05* lp4 17.0 8.00 0.47 lm1 16.4 10.0 0.60 PUPC 83/277* rm2 18.4 11.0 0.59 M. gluten AMNH 29862 P4 9.50 13.0 1.36 M1 13.0 15.0 1.15 M2 16.0 18.0 1.12 M3 16.0 16.0 1.00 AMNH 19993 m1 14.5 9.00 0.62 m2 16.5 11.0 0.66
Comparison
The studied material comprises the upper and the lower dentitions of medium size bovid.
The morphology of these specimens is typical of Miocene boselaphines in general; the divergent styles of the teeth make their inclusion in boselaphines. There are many
differences among the boselaphines from the Dhok Pathan. Selenoportax and
Pachyportax are large size boselaphines found in the Dhok Pathan Formation (Khan et al., 2007; 2009a). The Helicoportax, Elachistocerus and Eotragus are comparatively small size boselaphines (Pilgrim, 1937, 1939; Akhtar, 1992; Khan et al., 2009a). The medium size boselaphines include Tragoportax and Miotragocerus (Khan et al., 2009a).
The studied premolars and molars are well accentuated distinguished than those of
Tragoportax (Spassov and Geraads, 2004). Furthermore, the P2 is longer than the P3-P4, the P3 poorly expanded lingually and the metaconid of the p3-p4 is weak. The 51 morphology of the maxillar teeth and the mandibular teeth show that the samples reflects the diagnostic features of Miotragocerus and differentiate them to Tragoportax, other
Siwalik medium size boselaphine of the common stratigraphic range.
The morphology of the Dhok Pathan material allows it to be referred to Miotragocerus but the material metrically, is large in size than the Siwalik M. gluten (Table 5; Fig. 13).
Therefore, I suggest referring the Dhok Pathan material to M. large sp., while in the absence of horn-cores and cranial material species identification is impossible. However, the sample collected from the Dhok Pathan type locality is larger than the already known sample of Miotragocerus (Table 5). 52
Figure 11. Miotragocerus large sp. 1. PUPC 83/209 left maxillary ramus with P2-M1. 2.
PUPC 87/246 left maxillary ramus with DP3-M2. 3. PUPC 09/86 left mandibular ramus with p4-m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
53
Miotragocerus cf. gluten (Pilgrim, 1937)
Type Specimen: Skull lacking most of the face and the dentition: AMNH 19746.
Abbreviated Diagnosis (Pilgrim, 1937, 1939; Solounias, 1981; Spassov and Geraads,
2004): The horn-core is rather short or of moderate length, slender, slightly twisted, laterally compressed, situated immediately above the orbits and far apart on the frontals
with moderately prominent antero-internal and postero-external keels with flattened
lateral and slightly convex medial surfaces. The section is sub-elliptic. Anterior rugosities
at base of horn-cores usually strong, extending onto the frontal along the keel, which
often has several demarcations (steps) along its course. In front view, due to a slight
torsion of the horn-cores bases, the keels are often slightly convergent upwards in the
basal portion, then diverge towards the tips.
Studied Material: Horn-core: PUPC 01/01, incomplete, left side.
Description
Horn-core
The horn-core is broken at the apex. The horn-core is short, stout, slightly inclined backwards and compressed transversally (Fig. 12(1)). The antero-posterior diameter
(DAP) is greater than the medio-lateral diameter (DT). The horn-core has a weak clockwise twist. The horn-core has prominent anterior keel and weak posterior keel.
Between the keels the lateral surface is flat and the medial one is convex. The demarcations are well represented along the anterior keel. The anterior keel between the demarcations is well marked. The anterior keel torsion is weak and it is slightly convergent upwards at base then diverges towards the tips. The basal cross section is sub 54
triangular. The horn-core’s surface is without longitudinal grooves but the rugosities on
the anterior keel are common. The transverse diameter at the base is 21 mm and antero-
posterior diameter at the base is 30 mm. The mean index DT × 100/DAP is 70% at the
base. The length of the studied horn-core is ca. 113 mm. The transverse diameter of the
type AMNH 19746 at the base is 24.5 mm and its antero-posterior diameter at the base is
37 mm. The mean index DT × 100/DAP of the type is 66.2% at the base (Pilgrim, 1937).
Figure 12. Miotragocerus cf. gluten. 1. PUPC 01/01 left horn core: a) anterior view, b) medial view, c) lateral view, d) posterior view. Scale bar equals 30 mm total.
55
Comparison
Demarcations (steps) on the anterior keel are found in Miotragocerus and Tragoportax
(Spassov and Geraads, 2004). The horn-cores of Tragoportax are long and slender, usually curved backwards, with a triangular to subtriangular cross-section, well marked posterolateral keel and flattened lateral sides, but are less compressed than in
Miotragocerus. Anterior rugosities growing downwards from the anterior keel at the basis of horn-cores are absent or weak, and usually do not extend onto the frontal. Horn- cores have a heteronymous torsion (anti-clockwise on the right horn), so that the anterior
keels first diverge in anterior view, but they re-approach towards the tips.
The studied horn-core is laterally compressed, small, clockwise torsion, weak
posterolateral keel and clearly demarcated (Fig. 12(1)). All the characteristics are
diagnostic enough to include the sample in genus Miotragocerus. The horn-core cross section is elliptical. In lateral view the posterior face of the horn-core is appears flat, whereas the anterior one at the base becomes concave at the upper half, following an abrupt reduction of the horn-core dimensions towards the apex. The general morphology of this horn-core is clearly similar that of Miotragocerus gluten described from the
Middle Siwaliks (Pilgrim, 1937, 1939). Therefore, I provisionally assign the horn-core to
Miotragocerus gluten.
Discussion
The type species of Miotragocerus M. monacensis Stromer, 1928, was described from a partial skull from the Astaracian/Vallesian of Oberföhring. The genus Miotragocerus is known from several Upper Miocene localities of Europe (Solounias, 1981; Spassov and 56
Geraads, 2004) and the middle Pliocene of the Siwaliks in south Asia (Pilgrim, 1937;
1939; Akhtar, 1992). It is also known from Africa (Thomas, 1979). Solounias (1981) reviewed the genus, redescribed the species and referred Tragocerus valenciennesi to
Miotragocerus. Pilgrim (1937) named a genus Strepsiportax from Middle Miocene of the
Siwaliks which was synanomized to Protragocerus by Gentry (1970). Later on,
Solounias (1981) included Protragocerus gluten (Gentry) into Miotragocerus. Akhtar
(1992) reconfirmed the synonymy and added a new species Miotragocerus
dhokpathanensis Akhtar, 1992 from the Dhok Pathan. The new species was based on a
skull (PUPC 86/204) lacking most of the facial part, horn-cores and dentitions. The skull
is actually misidentified and it reflects the characteristics of Tragoportax. Consequently, it includes Tragoportax browni in this study after studying the metromorphological characteristics (see above).
Spassov and Geraads (2004) provided a comprehensive taxonomic revision of genera
Tragoportax and Miotragocerus, based on the materials collected from many Eurasian localities and noted that the differences in the literature are mostly metrical. They divided
Miotragocerus-Tragocerus-Tragoportax complex into two genera Miotragocerus and
Tragoportax as proposed by Solounias (1981). Solounias (1981) provided criteria for distinguishing between these two boselaphine groups. However, they are not confident of the separation criterion of Solounias (1981). The main differences between these two genera concern the horn-core pattern, some cranial and dental features and the presence of horns in females. Nevertheless, opinions among authors vary significantly and thus several species appear to interchange between the two genera (see discussions in Kretzoi 57
1968; Thomas 1979; Solounias 1981; Moyà-Solà 1983; Bouvrain & Bonis 1983;
Bouvrain 1988, 1994; Spassov & Geraads 2004).
The differences between Tragoportax and Miotragocerus have not always been clear and workers have differed over how best to diagnose them. Nevertheless, many workers proposed comprehensive study of the original material recently. According to them, many characters are misinterpreted and seem to be insufficient to categorize the
Miotragocerus-Tragoportax complex (e.g. see Gentry, 1971; Kostopoulos, 2005).
Two species of Miotragocerus, M. large sp. and M. gluten have been found in the Dhok
Pathan type locality. Miotragocerus large sp. is larger than that of M. gluten (Fig. 13) and only known presently by dental sample. Therefore, more perfect material is required for the specific identification.
58
M. large sp. M.gluten
P4 M1 16 20 14 18 12 16 10 14 12 8 10 Width
6 Width 8 4 6 2 4 0 2 0 0 2 4 6 8 10 12 14 0 2 4 6 8 10121416182022 Le ngth Length
M2 m1 22 14 20 18 12 16 10 14 12 8 10 Width Width 6 8 6 4 4 2 2 0 0 0 2 4 6 8 10 12 14 16 18 20 22 24 2 4 6 8 1012141618 Length Length
m2 12 10
8
6 Width 4
2
0 2 4 6 8 10 12 14 16 18 20 Length
Figure 13. Scatter diagram showing dental proportions of the Dhok Patan Miotragocerus.
Referred data are taken from Pilgrim (1937).
59
Genus PACHYPORTAX Pilgrim, 1937
Type Species: Cervus latidens (Lydekker, 1876).
Generic Diagnosis (Pilgrim, 1937): Boselaphinae of small to large or very large size;
closely allied to Strepsiportax but differing from that genus by the much more massive skull, with horn-cores longer, stouter, more twisted and less curved inwardly; occipital condyles and foramen magnum larger; mastoid process and squamosal shelf more developed; supraoccipital exposed on the upper surface of the occiput as a narrowly elliptical area much extended transversely; basioccipital approaching a rectangular shape, with posterior tuberosities not greatly expanded; upper molars strongly hypsodont but less so than in Selenoportax, quaderate, with strong entostyle, external folds weaker and less divergent than in Selenoportax, external ribs weaker than in Selenoportax, in
particular the median rib of the posterior lobe flattened, enamel rather thick, somewhat
less rugose than in Selenoportax, with traces of cement.
Stratigraphic Range: Middle Siwaliks.
Geographic Distribution: South Asia and Arabian Peninsula (Pilgrim, 1937, 1939;
Gentry, 1999; Khan et al., 2009a).
Pachyportax latidens (Lydekker, 1876)
Type Specimen: An isolated M3: GSI B219.
Diagnosis (Pilgrim, 1937, 1939; Gentry, 1999; Khan et al., 2009a): Horn-cores are
slender and situated directly above the orbits. They are stout with large size. They are less
divergence and twisted clockwise (right horn-core). The cross section is subtriangular. 60
The antero-internal and the postero-external keels are present. The third angle of the horn-cores is sufficient rounded to form a semicircle on the inner side of the keel.
A very large sized Pachyportax, with quadrate upper molars and strong entostyle much extended transversely which through its close association with the wings of the inner lobes produces one or more fossettes in the centre of the tooth; the crown is not constricted at the apex, relatively strong styles and ribs, enamel moderately thick and rugose with traces of cement. Crown is narrow at the base and broad at the apex in
Selenoportax whereas in Pachyportax the crown is not constricted at the apex. Entostyle is strong and much extending transversely in Pachyportax while in Selenoportax it is not much extending transversely. In Pachyportax posterior median rib is flattened whereas in
Selenoportax it is strong like anterior median rib.
Studied Material: Horn-core: PUPC 96/18, basal fragment, right side. Upper dentition:
PUPC 80/646, isolated M2. PUPC 80/203, isolated right M2. PUPC 09/74, isolated M2.
Lower dentition: PUPC 08/87, right mandibular ramus with m3 with broken hypoconulid and an anterior part of m2.
Description
The described material includes a horn-core (Fig. 14(1)), three upper molars and a mandibular fragment (Fig. 15(2-5)). The horn-core and the molars are partially damaged and somewhat deformed.
Horn-core
The anterior side of the horn-core pedicel is preserved and it is very well rugose. The dorsal orbital rim and the surface of the orbit are also preserved (Fig. 14(1)). The horn- 61
core’s inclination is very low in side view and it is less divergent. The cross section of the
horn-core is subtriangular. The lateral side is slightly convex. The medial surface is
somewhat rounded. The level of maximum transverse thickness lies posteriorly at the
base. The surface of the horn-core is furnished with numerous longitudinal furrows; nevertheless a deep longitudinal ridge running along the centre of the anterior side is
present. Both antero-internal and postero-external keels are present. The third angle does
not form a keel but is sufficiently rounded to form a semicircle on the inner side of the
keel axis. The length of the horn-core fragment is 92.5 mm. The anteroposterior (DAP)
and transverse basal diameters (DT) would be 50 × 43 mm, giving it an index of
compression at 86%.
Upper dentition
The cingulum is absolutely absent and all principal cusps the protocone, the paracone, the
metacone and the hypocone are present. The enamel is fairly thick and rugose. The
dentine is visible due to the early stage of wear. The entostyle is slightly damaged in
PUPC 80/646 and extended transversely. The fossettes are broad and filled with sand
stone (Fig. 15(2-4)). The posterior part of the anterior fossette is wider than the anterior
part. The entostyle of PUPC 80/203 is badly damaged and its small portion can be seen at
the base of the transverse valley of the molar. The styles are well developed; the
mesostyle is strongly developed while the metastyle is moderately developed. The
postprotocrista is clearly at the anterior end of the hypocone. The molars are
quadrangular in shape (Fig. 15(2-4)). A prominent and distinct pressure mark lies at the
anterior and the posterior sides of the molars verifying their second position in the molar
series. 62
Lower Dentition
It is a hypsodont and narrow crowned specimen (Fig. 15(5)). The metaconid and entoconid are slightly damaged at the apex while all ribs and stylids are well developed.
The average thickness of the enamel all over the crown is 2.0 mm and it is smooth lingually but finely rugose on the labial side. The cement is present lingually. The ectostylids are well developed lying towards the hypoconid with oval cross section (Fig.
15(5)). The metaconid is spindle shaped with an inflated median part, which produces the moderately strong anterior median rib of the metaconid. The anterior transverse flange is very prominent in the molars. The hypoconid is more constricted than those of the protoconid. The exact measurement of the specimen is not possible owing to the broken parts of the teeth (Table 6).
63
Table 6: Comparative measurements of the cheek teeth of P. latidens (Lydekker) in mm
(millimeters). * The studied specimens. Referred data are taken from Pilgrim (1937,
1939), Akhtar (1992), Khan et al. (2009a).
Taxa Number Nature/Position Length Width W/L ratio P. latidens PUPC 80/646* ?M2 31.0 ca. 19.0 0.61 PUPC 80/203* rM2 27.0 ca. 18.5 0.68 PUPC 09/74* ?M2 25.0 20.0 0.80 PUPC 98/59 rM2 22.0 17.3 0.78 PUPC 96/40 rM2 19.4 18.4 0.94 PUPC 96/3 rM2 27.0 22.0 0.81 PUPC 86/37 rM2 27.4 18.0 0.65 PUPC 86/36 rM2 30.0 23.0 0.76 PUPC 83/718 rM2 27.4 26.0 0.94 PUPC 83/646 rM2 30.0 18.0 0.6 PUPC 83/744 rM2 30.2 21.9 0.72 PUPC 86/210 rM2 26.0 17.1 0.65 PUPC 00/100 rM2 25.5 25.0 0.98 PUPC 04/14 rM2 29.3 20.6 0.70 PUPC 98/60 rM2 23.1 15.9 0.68 PUPC 97/103 rM2 24.5 17.7 0.72 PUPC 86/203 rM2 26.4 17.9 0.67 AMNH 29964 rM2 28.0 25.0 0.89 AMNH 19730 rM2 28.5 28.5 1.00 P. nagrii PUPC 99/50* ?M3 19.5 18 0.98 GSI B808 M3 21 17.2 0.81
64
Comparison
The probable large size of the horn-core, wide insertion and the position of the keels agree with the large Siwalik boselaphines Selenoportax and Pachyportax (Pilgrim, 1937).
It agrees with P. latidens rather than S. vexillarius in a very low inclination of the horn- core in side view, outer face between the keels slightly convex and rounded, third angle is internal and distinct, and less divergence of the horn-core. It could also be similar to the horn-core of P. latidens (Lydekker) var. dhokpathanensis from the Dhok Pathan
Formation of the Middle Siwaliks, GSI B488, referred by Pilgrim (1939: 200) and to the left horn-core of a large boselaphine or bovine from Piram Island, BMNH M2402a, referred by Pilgrim (1937: 746; 1939: 175). The horn-cores of Parabos cordieri from the
French early Pliocene were similar to those of Pachyportax.
The holotype GSI B488 of P. latidens (Lydekker) var. dhokpathanensis has DAP and DT of 56 × 42 mm, giving it an index of compression of 75% and the referred skull GSI
B245 of P. latidens (Lydekker) var. dhokpathanensis has DAP and DT of 62 × 49 mm, giving it an index of compression of 79% (Pilgrim, 1939) and the holotype PUPC 87/323 of P. giganteus (Akhtar, 1992) has DAP and DT of 78 × 52 mm, giving it an index of compression of 66.6%. Akhtar (1992) described two horn-cores PUPC 85/103 and PUPC
85/104 (formerly PMNH) from the Hasnot of the Dhok Pathan Formation. PUPC 85/103 has DAP and DT of 62 × 48 mm, giving it an index of compression of 77.4% and PUPC
85/104 has DAP and DT of 59 × 45 mm, giving it an index of compression of 76.2%
(Akhtar, 1992). The compression index of the studied horn-core is 86%, indicating a
stouter sample than those of the earlier studied specimens (Fig. 14(1)). 65
The teeth are hypsodont and have large transversely expanded entostyles (Fig. 15(2-5)).
They have moderate labial folds and the median ribs are narrow. The posterior median rib is rather less prominent. The teeth differs from a giraffids in that the teeth are too hypsodont, their entostyles are too large and the folds are strongly compressed. They are definitely from a boselaphine bovid of a bovine stamp on account of the anteroposterior compression of the lobes, the strong inner folds of the outer lobes, the prominence and transverse extension of the entostyles but the weak median ribs, and the stratigrphic level distinguish them from the bovine (Pilgrim, 1939; Akhtar, 1992; Khan et al., 2009a). They are of appropriate size and morphometrically to match the holotype of P. latidens (Table
6). The studied sample is sufficient for the specific determination and consequently P. latidens is assigned for the sample. 66
Figure 14. Pachyportax latidens. 1. PUPC 96/18 right horn core: a) anterior view, b) medial view, c) posterior view, d) lateral view. Scale bar equals 30 mm total.
67
Figure 15. Pachyportax latidens. 2. PUPC 80/646 isolated M2, side indeterminate. 3.
PUPC 09/74 isolated M2, side indeterminate. 4. PUPC 80/203 isolated right M2. 5.
PUPC 08/87 right mandibular ramus with m3 with broken hypoconulid and an anterior part of m2. 6. Pachyportax cf. nagrii. PUPC 99/50 isolated molar, side indeterminate.
Occlusal views. Scale bar equals 30 mm total.
Pachyportax cf. nagrii
Studied Material: Upper dentition: PUPC 99/50, partially damaged isolated M3.
Description and Comparison
The molar is probably third in molar series because the metastyle is pillar like structure and the posterior facet is not contact (Fig. 15(6)). The protocone of the molar is broken away. The anteroposterior length of the anterior lobes is less than that of anteroposterior 68
length of the posterior lobes (Table 6). The styles are less prominent than Selenoportax
and Pachyportax latidens. The crown neck is not narrow as in Selenoportax (Khan et al.,
2009a). The entostyle is expanded transversely. The lingual lobes are less constricted and
the cavities are open. The molar is smaller in size than P. latidens (Fig. 16; Table 6) and
nevertheless it reflects the characteristics of Pachyportax. Lacking adequate material, it is referred to as P. cf. nagrii.
Discussion
Pilgrim (1937) erected the genus Pachyportax when he applied the generic name
Pachyportax to all the specimens, which were described and figured by Lydekker in 1876 under the name Cervus latidens. Pilgrim (1937, 1939) mentioned two species P. latidens
Lydekker and P. nagrii, one subspecies P. latidens dhokpathanensis and one variety P. latidens var. dhokpathanensis from the Middle Siwaliks. According to Pilgrim (1939) P. nagrii occurred earlier in the Siwalik layers. Akhtar (1995) erected a new species P. giganteus and synonymized the subspecies P. latidens dhokpathanensis with the species
P. latidens (Lydekker) Pilgrim. The genus was originally founded to incorporate the large boselaphines of the Siwaliks, which probably have a closer ancestry to the Bovini than
Tragoportax (Gentry, 1999).
Pachyportax is considered as a typical Late Miocene taxon and occurs in the Nagri and the Dhok Pathan formations of the Siwaliks (Fig. 16), until Proamphibos replaces it soon
after the start of the Tatrot (Khan et al., 2009a). A date around 7.0 Ma, during the Latest
Miocene (Barry et al., 1991) would be considered as a possible date for a fauna containing P. latidens. The type species originated in the Late Miocene, possibly 69
extending its range into the Pliocene. The type specimen of P. latidens is an upper molar,
but it is best represented by a cranium described by Pilgrim in 1939 and stored in the
Calcutta Museum (India), a cast of which is found in London (BMNH), BMNH M26573.
A very similar boselaphine to Pachyportax is Parabos Arambourg and Piveteau of which
the type and only species, Pachyportax or Parabos cordieri, comes from the Lower
Pliocene of Montpellier, southern France. According to Gromolard’s revision (1980) of
Parabos, it may be assumed that this generic name is the senior synonym for
Pachyportax. Around the end of the Late Miocene boselaphines gave way to bovini or
bovine like bovids (Gentry, 1999).
M3 P. latidens P. nagrii 40 35 30 25 20
Width 15 10 5 0 0 5 10 15 20 25 30 35 40 Length
Figure 16. Scatter diagram showing dental proportions of the Dhok Pathan Pachyportax.
Referred data are taken from Pilgrim (1937, 1939), Akhtar (1992) and Khan et al.
(2009a).
70
Genus SELENOPORTAX Pilgrim, 1937
Type Species: Selenoportax vexillarius Pilgrim, 1937.
Generic Diagnosis (Pilgrim, 1937, 1939; Akhtar, 1992; Khan et al., 2008a): Moderate
to large sized Siwalik bovid; with hypsodont to extremely hypsodont teeth, upper molars
quadrate with strong divergent styles, median ribs well developed, entostyle strongly
developed and ectostylid moderately developed, enamel very rugose (Pilgrim, 1937).
Crown is narrow at the base and broad at the apex in Selenoportax whereas in
Pachyportax the crown is not constricted at the apex. Entostyle is strong and much
extending transversely in Pachyportax while in Selenoportax it is not much extending
transversely (Khan et al., 2008a). In Pachyportax posterior median rib is flattened
whereas in Selenoportax it is strong like anterior median rib (Akhtar, 1992).
Stratigraphic Range: Middle and Upper Siwaliks (Akhtar, 1992; Khan et al., 2009a).
Geographic Distribution: South Asia (Pilgrim, 1937, 1939; Akhtar, 1992; Bibi, 2007;
Khan et al., 2009a).
Selenoportax cf. vexillarius Pilgrim, 1937
Type Specimen: A skull lacking maxilla and dentition and most of the basicranium
(AMNH 19748).
Abbreviated Diagnosis (Khan et al., 2008a, 2009a): Cheek teeth large and strongly
hypsodont, enamel very rugose. Upper molars quadrate with strong and divergent styles
near the neck of crown, ribs quite large, entostyle/ectostylid strongly developed. Fossettes
without indentations and simple in outline, anterior transverse flange poorly developed at 71 front of lower molars. S. vexillarius is smaller than that of S. lydekkeri which was the large one species of the Siwalik Selenoportax.
Studied Material: Lower dentition: PUPC 99/47, right mandibular ramus with m2 and roots of m1. PUPC 99/22, isolated left semierupted molar. PUPC 00/45, isolated left m2.
PUPC 07/16, partially damaged molar. PUPC 97/02, isolated right m3. PUPC 97/01, isolated right m3.
Description
The sample of Selenoportax comprises only lower dentition (Fig. 17(1-3)). Three out of six are preserved with all morphological features whereas rest of the three is partially damaged. A mandibular ramus having length 73 mm bears m2.
Lower dentition
The specimens of the lower dentition present well developed second and third molars
(Fig. 17(1-3); Table 7). The molars have fully developed principal conids. The conids are narrower than the cones and the anterior transverse flange is moderately developed in the lower molars. The molars present various stages of wear and have prominent ectostylids.
In the third molars the hypoconulid is well developed, long and narrow, with a wide and inflated central area having two sloping cristids. The ribs and stylids are divergent lingually.
72
Table 7: Comparative measurements of the cheek teeth of S. cf. vexillarius in mm (millimeters). * The studied specimens. Referred data are taken from Pilgrim (1937, 1939), Akhtar (1992) and Khan et al. (2009a).
Number Nature/Position Length Width W/L S. cf. vexillarius PUPC 99/47* rm2 23.0 12.0 0.52 PUPC 99/22* lm? 26.5 - - PUPC 00/45* lm2 26.0 13.4 0.51 PUPC 07/16* m? 25.4 13.0 0.51 PUPC 97/02* rm3 27.0 14.0 0.51 PUPC 97/01* rm3 28.0 13.0 0.46 S. vexillarius PUPC 04/1 lm2 27.9 16.1 0.57 lm3 31.4 16.0 0.50 PUPC 98/78 lm2 25.0 16.0 0.64 lm3 36.0 15.0 0.41 PUPC 85/40 lm1 19.7 12.5 0.63 PUPC 04/12 lm2 20.0 12.5 0.62 PUPC 87/90 lm3 38.0 16.5 0.43 AMNH 10514 lm3 33.0 15.0 0.45 AMNH 29917 lm1 18.0 13 0.72 AMNH 19844 lm2 25.9 16.5 0.63 AMNH 19514 lm2 22.0 15.5 0.70 AMNH 29917 lm2 21.0 15.0 0.71 AMNH 19514 lm3 33.0 21.5 0.65
Comparison
The specimens are large enough to include them the large Siwalik boselaphines of the late Miocene Selenoportax and Pachyportax (Pilgrim, 1937; Khan et al., 2009a). The general contour of the studied specimens, the rugosity of the enamel, the strong ectostylids, the prominent median ribs, constricted labial lobes, the strong and divergent 73
stylids (Fig. 17(1-3)) exclude the specimens from the genus Pachyportax and favor their
inclusion in the genus Selenoportax. Selenoportax is represented by two species in the
Siwaliks S. vexillarius and S. lydekkeri. The dimensions and the morphology of the
studied material (Fig. 18) reveal all the features of the species S. vexillarius as cited by
Pilgrim in 1937 and 1939 (Table 7). The studied specimens present the same
morphological features of the holotype (Pilgrim, 1937). The paraconid, the parastylid, the protoconid, the entoconid, and the development of the stylids are fairly similar. The
second molars are narrow crowned, as indicated by its width and length ratio, and
resemble with the specimens AMNH 19844, AMNH 19514 and AMNH 29917 as
discussed by Pilgrim in 1937. The enamel layer of the third molar is finely plicated and it
compares very favorably with AMNH 19514 (Pilgrim, 1937). The studied sample
morphometrically (Fig. 18; Table 7) resembles with S. vexillarius, however, the material
only comprises lower dentition and for the exact specific determination more material is
required. Therefore, S. cf. vexillarius is assigned for the lower dentition.
74
Figure 17. Selenoportax cf. vexillarius. 1. PUPC 99/47 right mandibular ramus with m2 and roots of m1. 2. PUPC 97/02 isolated right m3. 3. PUPC 97/01, isolated right m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
75
Discussion
Pilgrim (1937) described two species of genus Selenoportax, S. vexillarius and S.
lydekkeri based on a collection from various Siwalik localities in Pakistan and India.
Akhtar (1992) ascribed both S. vexillarius and S. lydekkeri from the Middle and Upper
Siwaliks in the genus Selenoportax and added two new species in it, S. dhokpathanensis
and S. tatrotensis; the former was based on a damaged cranium (PUPC 86/248) and the
later was based upon a maxillary portion bearing right P3-M3 and left P3-M3 (PUPC
87/19). More recently, Khan et al. (2009a) synanomyzed S. dhokpathanensis with S.
lydekkeri and S. tatrotensis with S. vexillarius. Therefore, the genus Selenoportax
consists only of two valid species and these are S. vexillarius and S. lydekkeri. S.
vexillarius is smaller in size than S. lydekkeri (Fig. 18). In general the cheek teeth of S.
vexillarius are smaller than those of S. lydekkeri (Fig. 18).
Selenoportax records from the late middle Miocene and Pliocene of the Siwaliks (Akhtar,
1992; Barry et al., 2002; Bibi, 2007), nevertheless some uncertain occurrences are
mentioned out of the Siwaliks (Solounias, 1981). The specimen collected from the Tatrot
formation (Akhtar, 1992) extends the range of Selenoportax from 10.3 to 2 Ma, contrary
to the findings of Barry et al. (2002) who stated a maximum range of 10.3-7.9 Ma for this
genus. Solounias (1981) referred a Selenoportax horn core from Pikermi; according to
him the material is not diagnostic enough for a more specific determination. So, this
supposed occurrence of a single horn core cannot be taken as indicative of a Selenoportax
presence in Europe. Qiu and Qiu (1995) list Selenoportax sp. from the Lufeng fauna
(Chinese Miocene), age of 11.1-8.0 Ma (Steininger, 1999; Flynn and Qi, 1982). These 76
specimens represent the sole record of Selenoportax from the Miocene of northern and
central Asia (Tedford et al., 1991; Nakaya, 1994; Qiu and Qiu, 1995).
S. cf. vexillarius S. lydekkeri S. vexillarius
m1 m2
20 20 18 18 16 16 14 14 12 12 10 10 Width 8 Width 8 6 6 4 4 2 2 0 0 0 5 10 15 20 25 30 0 5 10 15 20 25 30 35 Length Length
m3 24 22 20 18 16 14 12
Width 10 8 6 4 2 0 0 5 10 15 20 25 30 35 40 45 Length
Figure 18. Scatter diagram showing dental proportions of the Dhok Pathan Selenoportax. Referred data are taken from Pilgrim (1937, 1939), Akhtar (1992) and Khan et al. (2009a).
77
Subfamily ANTILOPINAE Gray, 1821
Tribe ANTILOPINI Gray, 1821
Genus PROSTREPSICEROS Major, 1891
Type Species: Prostrepsiceros houtumschindleri (Rodler and Weithofer, 1890).
Generic Diagnosis (Bibi and Gulec, 2008): Horn-cores with anticlockwise torsion, open spiraling, amounting to about 180° total torsion (half a rotation); moderately strongly divergent; inserted above orbit; mediolaterally compressed; cross-section ovate to kidney-shaped, with lateral surface flat basally to slightly concave distally; posterolateral keel present; anterior keel variably present, always less sharp than posterolateral keel; surfaces with longitudinal grooving present; horn-cores extending onto pedicels anteriorly. Supraorbital foramina small, circular, opening anteriorly, and placed in semi- triangular pit. Post-cornual fossa present, wide and deep. Frontal sinuses small, not extending into pedicel. The p3 bearing separate paraconid parastylid; backwardly-slanted metaconid that fuses with entoconid in early wear stages; large open lingual valley between paraconid and metaconid; entoconid and entostylid well separated but fusing in mid-wear stages; entoconid much larger than entostylid; hypoconid weak; some cementum present in the lingual valleys. Upper teeth differ from those of Gazella in molars of larger size; with metacone more rounded; wider paracone rib; mesostyle wider and more projecting; and metastyle wider.
Stratigraphic Range: Lower and Middle Siwaliks. 78
Geographic Distribution: From Middle Miocene to Late Miocene. Prostrepsiceros is a
bovid common in the late Miocene Europe, Africa and the Subcontinent (Gentry, 1971;
Solounias, 1981; Bouvrain, 1982; Kostopoulos and Koufos, 1996; Pilgrim, 1939).
Prostrepsiceros cf. vinayaki (Pilgrim, 1939)
Type Specimen: Left horn-core: GSI B799.
Emended Diagnosis (modified from Pilgrim, 1939): Bovidae of small size with horn-
cores forming an open clockwise spiral and torsioned, with an elliptical to ovate cross
section. Two keels are present, a variable anterior keel and sharp posterolateral keel. The
“anterior” one descends to the postero-medial side of the base, forming a complete gyre towards the apex. Thin discontinuous furrows run along their surfaces. A deep longitudinal furrow, starting above the base, runs in parallel with anterior keel.
Nevertheless, the presence and development of this furrow is variable. The horn-cores curve moderately backwards, whereas they are re-curved upwards at their top with
convex medial side and slightly flat lateral one.
Studied Material: Horn-cores: PUPC 87/319, almost complete, left side. PUPC 87/318,
incomplete, probably right side.
Description
The described material comprises only horn-cores. One specimen (PUPC 87/319) is
almost completely preserved and the second one (PUPC 87/318) is a fragment (Fig. 19(1-
2)).
79
Horn-cores
Horn-cores are mediolaterally compressed (Fig. 19(1-2)). The horn core’s cross section is symmetrical oval with convex medial side and slightly flat lateral one. In the uppermost part of the horn core (PUPC 87/319) the cross section becomes sub-triangular. Horn- cores with anticlockwise torsion and open spiraling curve moderately backwards, whereas they are re-curved upwards at their top. The degree of torsion with the twisted axis of the horn-core produces a general outward curvature in its lower half. The surface is longitudinally striated and there are deep furrows at the postero-external and the anterior corners. Nevertheless, the presence and development of the furrows are variable: these are very deep and double in PUPC 87/318 but less deep in PUPC 87/319. Two keels are present. The “anterior” one descends to the postero-medial side of the base, forming a complete gyre towards the apex. It is smooth in the lower part of the horn-core but becomes sharper in its upper half. A deep longitudinal furrow, starting 2-3 cm above the base, runs in parallel with this keel or even replaces it. The second keel descends over the anterior margin of the postcornual groove and also forms a complete gyre towards the top. The preserved length of PUPC 87/319 is 158 mm, the DAP is 33 at the base and the
DT is 26 mm at the base. The mean index DT × 100/DAP is 78% at the base.
Comparison
The general morphological characters of the available horn-cores allot the medium sized antelope of the Middle Siwaliks to Prostrepsiceros (Pilgrim, 1939; Thomas, 1984). The deep anterior and posterior furrows, extremely torsioned, ovate cross section, two keels and mediolaterally compressed (DAP > DT) observed in the studied sample places them 80
into Prostrepsiceros (Fig. 19(1-2)). The late Miocene Prostrepsiceros is poorly known
from the Dhok Pathan type locality (Pilgrim, 1939; Thomas, 1984; Akhtar, 1992;
Kostopoulos, 2005). In my knowledge the Dhok Pathan sample constitutes at present
time one of the best known evidence of this genus in the Middle Siwaliks of Pakistan.
Prostrepsiceros vinayaki (Pilgrim, 1939) is a known species of the Siwalik
Prostrepsiceros (Pilgrim, 1939; Gentry, 1999; Kostopoulos, 2005). The observed
characters in the studied sample are present in the Siwalik P. vinayaki and the material is
very good matches for P. vinayaki. It agrees with the Siwalik P. vinayaki in having deep anterior and posterior furrows, extremely torsioned, ovate cross section, two keels and mediolaterally compression. It is very similar to the holotype horn-core (GSI B799) of P. vinayaki in having almost same index of compression. The DAP and DT at the base of
GSI B799 being 25.5 × 19.5 mm, giving an index of compression of 76%. It is, however, slightly larger, in the studied sample, has index of compression 78%. Prostrepsiceros aff.
vinayaki of Emirates of Abu Dhabi has anteroposterior and transverse basal diameters of
29.6 × 20.9 mm, giving it an index of compression of 71%. Nisidorcas planicorni
(Pilgrim, 1939) of Peram Island and Graeco-Turkish Turolian (Bouvrain, 1979; Kohler,
1987) is similar sized, and compressed to about the same extent, but has a posterior keel
strengthening distally and no anterior keel (Gentry, 1999).
According to the available data from Eurasian several localities (Pilgrim 1939; Gentry
1999; Kostopoulos, 2005; Bibi and Gulec, 2008) P. vinayaki differs from Eurasian
Prostrepsiceros P. vallesiensis, P. houtumschindleri, P. rotundicornis, P. fraasi and P. zitteli in the longer and slenderer horn-cores, which are more torsioned than spiraled and extremely compressed mediolaterally (Bouvrain, 1982; Gentry, 1999). 81
Discussion
Pilgrim (1939, p. 42) created a new species Gazellinae (?) gen. indet. (aff. Helicotragus) vinayaki, based on a very fragmentary left horn-core. Gentry (1970, p. 259) supposed that the holotype left horn-core GSI B799 reflects characters of young Sivoreas eremite. The holotype was collected from the locality Nila of Dhok Pathan Formation. Thomas (1984) studied two more horn-coresYGSP 4082 and YGSP 2631, collected from the same level of the Dhok Pathan Formation and ascribed species ‘vinayaki’. He observed that the holotype and the new sample clearly distinguished from Sivoreas eremite. Having observed the close resemblance of the sample with Prostrepsiceros, Thomas (1984) allotted the material to P. vinayaki (Pilgrim, 1939). The studied horn-cores collected from the type locality Dhok Pathan morphometrically resemble with the previous sample and reflect the morphometric characteristics of P. vinayaki. I therefore consider the horn- cores to P. vinayaki and confirm here its presence in the type locality of the Dhok Pathan
Formation.
Prostrepsiceros occurs in a wide geographic area of Saudi Arabia, North Africa, Greece,
Tibetan Plateau and the Subcontinent (Pilgrim, 1939; Thomas, 1984; Bouvrain 1982;
Bouvrain and Thomas 1992; Gentry 1999, 2003). Prostrepsiceros has two or more lineages and is the most diversified genus. Some species have keels and inclined insertions of the horn-cores; others have no vestigial keels, open spiraling, more or upright insertions and little inclination of their cranial roofs. The radiation and distribution of the genus, as well as the phylogenetic relationships among the quite numerous affiliated taxa are not yet clear enough (Kostopoulos, 2009). The genus appeared in 10 Ma (Gentry, 2003) and exists hereafter with a variety of forms (Bouvrain 82 and Thomas, 1992; Kostopoulos and Koufos, 1996; Bonis and Koufos, 1999), suggesting a fast radiation and distribution.
Early representatives of Prostrepsiceros share in common a weak cranio-facial angle, a long basioccipital with a wide central groove, small supraorbital foramens not into pits and away from the frontal base, small postcornual grooves and slender horn-cores comparatively to the skull size, inserted on the posterior part of the orbital roof, strongly inclined backwards and widely separated on the frontals. Some of these apparently primitive features can also be seen in the poorly known Sinapodorcas from Middle Sinap,
Turkey (Bouvrain et al., 1994; Gentry, 2003), a genus that seems to me very closely related to Prostrepsiceros.
Bouvrain’s (1982) suggestion in dividing Prostrepsiceros in two distinct subgenera might indeed reflect an early division in two distinct lineages that evolved independendly from late Vallesian onwards. The first one includes P. vallesiensis, P. houtumschindleri, P. lybicus and P. vinayaki and the second Sinapodorcas, P. syridisi, P. rotundicornis, P. fraasi, P. zitteli and P. axiosi. According to Lehmann and Thomas (1987) a relationship between P. lybicus and the Jebel Hamrin form seems undeniable while Gentry (1999) looks for the possible ancestor stock of Sahabi form in P. fraasi from Samos.
Kostopoulos (2009) suggests a possible relation between P. lybicus, P. vinayaki and P. vallesiensis. According to Gentry et al. (1999) Antilope itself probably descends from within Prosterpsiceros. The ancestry of Antilope cervicapra, the living blackbuck of
India lies within Prostrepsiceros (Gentry, 1999). 83
Figure 19. Prostrepsiceros cf. vinayaki. 1. PUPC 87/319 left horn-core. 2. PUPC 87/318
?right horn-core. a) anterior view, b) medial view, c) lateral view, d) posterior view.
Scale bar equals 30 mm total. 84
Genus GAZELLA Blainville, 1816
Type Species: Gazella dorcas Linn., 1758.
Generic Diagnosis (Pilgrim, 1939; Kostopoulos, 2009): Upper molar hypsodont, with prominent narrow styles, without entostyles, fossettes crescentic in shape, median ribs moderately strong in primitive forms, weak or absent in progressive forms, P2 longer than P3 and P4. Lower molars with goat folds, small ectostylids, fossettes with fairly simple outline, stylids and ribs moderately developed. Horn-cores moderately long, spaced, inserted over the orbits, not very divergent, sub circular or elliptical in cross section, unkeeled, inward or forward curvature near the tip, surface covered by longitudinal striae.
Stratigraphic Range: Lower to Middle Siwaliks (Middle Miocene to Pliocene).
Geographic Distribution: The occurrence of Gazella is recorded from the Lower
Pliocene of Eurasia and several middle Miocene and Pleistocene localities of Africa
(Gentry, 1999). It is abundantly found in the Lower Pliocene fauna of Asia and the southern parts of Europe (Kostopoulos, 2009). It is recorded from the Siwaliks of the
Subcontinent (Pilgrim, 1937, 1939; Thomas, 1984; Akhtar, 1992; Khan, 2007).
Gazella lydekkeri Pilgrim, 1939
Type Specimen: A skull and conjoined mandible (AMNH 19663).
Diagnosis (Pilgrim, 1937, 1939): Upper molars moderately hypsodont, styles narrow and strong with entostyles very small or absent, enamel moderately thick and rugose, fossettes narrow and deep, anterior median rib stronger than posterior one, premolar series rather long. Lower molars extremely hypsodont approaching quadrate shape, with 85 small ectostylids, prominent goat folds, fossettes fairly simple in outline, stylids and ribs moderately developed. Horn cores moderately long, spaced, slightly curved backward, broadly elliptical in cross-section, fine ribs becoming rudimentary near the tips, one deep furrow posteriorly.
Studied Material: Horn-cores: PUPC 83/319, almost complete with a piece of frontal, broken at the apex, left side. PC-GCUF 09/45, almost complete with supraorbital pit, left side. Lower dentition: PUPC 08/136, right isolated m2. PUPC 07/13 and PUPC 09/76, isolated left m3s.
Description
The material comprises two horn-cores, a second lower molar and two lower third molars
(Fig. 20).
Horn-cores
The horn cores curve gently upwards and backwards (Fig. 20(1-2)). The horn core looks to diverge very slightly and taper towards the tip. The supraorbital pit is preserved in PC-
GCUF 09/45. It is pear shaped. The horn cores are relatively short, smart and present directly above the orbit. The bases of the horn cores are strongly elliptical, without keels, untwisted and slightly more convex on the inner side than on the outer side. The surfaces of the horn cores are furnished by the longitudinal furrows and there is one deep furrow present posteriorly. The horn cores arise close together and diverging gradually. The horn cores are supported by the smooth pedicels. The horn cores look alike morphologically and metrically. The preserved length of PUPC 83/319 is 72.6 mm, the DAP is 23.5 mm at the base and the DT is 18.5 mm at the base. The mean index DT × 100/DAP is 78.7% at 86 the base. For PC-GCUF 09/45 the length is 60.5 mm, the DAP is 21.5 mm, the DT is
17.5 giving the index of compression is 81.3%.
Lower dentition
The m2 is well preserved and subhypsodont (Fig. 20(3)). The ribs are broader at the base and gradually narrow toward the tips. The anterior median rib is more evident than the posterior one. The mesostylid on the lingual side is very weak, visible only on the upper half of the tooth crown. On the lingual side, a short tubercle is present at the base between the metaconid and the entoconid. The labial transverse valley is occupied at the base by a short and robust ectostylid. A prominent goat fold is present along the anterior surface of the tooth. All the cusps are well preserved and conical in form. The lingual cusps are higher than the labial ones. The fossettes are moderately wide and simple. The metaconid is slightly higher than the entoconid.
PUPC 07/13 and PUPC 09/76 are well preserved left isolated third lower molars (Fig.
20(4-5)). The enamel is rugose especially on the lingual side. All the conids are well preserved and the lingual ones are pointed. The anterior goat fold is broad and prominent, and the hypoconulid is narrow and crescentic (Fig. 20(3-5)). The ectostylid is robust, rounded and relatively longer. The mesostylid is narrow, gradually prominent towards the apex. The comparative measurements are provided in table 8.
87
Table 8: Comparative measurements (mm) of the cheek teeth of G. lydekkeri. * The studied specimens. Referred data are taken from Pilgrim (1937, 1939), Akhtar (1992) and Khan (2008). Number Nature/Position Length Width W/L ratio PUPC 84/67 rm2 16.6 10.0 0.60
PUPC 87/160 lm2 14.0 9.00 0.64
PUPC 04/02* rm2 14.0 8.70 0.62
rm3 19.0 8.60 0.45
PC-GCUF 09/16 lm2 16.0 11.0 0.68 lm3 21.0 10.5 0.5 PUPC 04/08 lm3 20.0 9.00 0.45 PUPC 86/04 lm2 15.0 10.0 0.66
PUPC 87/ 162 lm3 22.0 10.5 0.47
PUPC 67/42 rm3 20.0 9.00 0.45 PUPC 08/136* rm2 15.5 7.50 0.48 PUPC 07/13* lm3 22.0 9.50 0.43 PUPC 0976* lm3 23.7 10.6 0.44
Comparison
The horn cores of Gazella are rising almost directly above the orbits and gently curving
upwards and backwards. The horn cores characterizing an oval cross section with
mediolateral compression are indicative of Gazella (Kostopoulos, 2009, Pilgrim, 1937,
1939; Akhtar, 1992). The studied horn cores reflect similar morphological characters with G. lydekkeri characteristically arises immediately above the orbit, moderated length, without keel, less mediolaterally compression, small but deep lachrymal fossa and has elliptical cross section (Fig. 20; Table 8). The horn cores refer to G. lydekkeri while 88
noting its morphometric characteristics. The small size and the less laterally compression
of the horn-cores is somewhat distinguish it to male G. lydekkeri and indicates that the horn-cores belong to female animal (Gentry, 1970).
The small and deep lachrymal fossa is present in G. lydekkeri as in living G. granti and very different from the large and deep fossa in living G. thomsoni (Bibi and Gulec, 2008).
The horn cores differ from those of G. capricornis because of not having circular cross section. It is different from G. pilgrimi which has almost straight horn cores with more persist few longitudinal ribs (Kostopoulos, 2005). Gazella deperdita also has more curved horn cores. Gazella mytilinii is a large species with narrowly elliptical, almost parallel and little curved horn cores (Kostopoulos, 2009). The horn core differs from G. dorcas, G. cuvieri and G. rufifrons by having less mediolateral compression. The horn cores of G. schlosseri are more slender and upright and the ribs extend to the tip. Gazella longicornis has exceptionally long horn cores. Gazella rodleri has spirally twisted horn cores. Gazella anglica and G. daviesii are small having highly compressed horn cores
(Pilgrim, 1937). The superorbital pit in the studied sample is shorter and less deep than in
G. subgutturosa. In G. capricornis the horn cores are elongated and strongly curved backwards as G. deperdita (Heintz, 1971). Gazella pilgrimi and G. gaudryi are small sized species (Kostopoulos, 2009). In G. pilgrimi the horn cores are strongly inclined backwards (Kostopoulos, 2009).
The studied dental material of G. lydekkeri recovered from the type locality of the Dhok
Pathan Formation from the Middle Siwaliks of Pakistan afford no justification for separating them generically from Gazella: small ectostylids and prominent goat folds on 89 the lower molars are the characters that support their inclusion into the genus Gazella
(Fig. 20-21).
Morphological and metrical characters of the dental specimens indicate that G. lydekkeri is similar in appearance and size to the living G. bennetti: longer and slenderer skull, higher occipital and the shape and direction of horn cores are very similar in both species
(e.g. see Pilgrim, 1937, 1939; Akhtar, 1992; Kostopoulos, 2009). However, there are some characters that differentiate the living G. bennetti from the extinct G. lydekkeri.
These are the acquisition of horns in females, lengthening of horn cores, greater bending down of face on cranial axis, shortening of nasals, increase in hypsodonty, reduction of premolar series and enlargement of bullae. Gazella lydekkeri differ from G. bondi (Cooke and Wells, 1951), G. paotehensis and G. gaudryi (Bohlin, 1941) in greater hypsodonty.
This suggests tougher diet for G. lydekkeri than the later two species. This assumption is supported by the results of Merceron et al. (2006) who suggested, by micro-wear analysis, abrasive grazing and browsing habits for the gazelles of Bulgaria. In G. dorcadoides and G. altidens the teeth are more hypsodont than in G. lydekkeri. Gazella lydekkeri is similar to G. dorcadoides in hypsodonty but is different due to equally strong anterior and posterior median ribs of the later. Gazella lydekkeri is smaller with longer premolars, shorter, slenderer and remarkably less compressed horn cores than G. pilgrimi and G. capricornis from Turkey (Kostopoulos, 2005). The length of post canine teeth series in G. lydekkeri is greater than that of G. capricornis. In both these species P2 is longer than P3, P3 than P4 and p3 than p4. Premolars of G. lydekkeri are, however, narrower than those of G. capricornis. Entostyles are most frequently found in M1 and
M2 of G. capricornis and less frequently in G. gaudryi, G. dorcadoides and G. lydekkeri. 90
Fairly strong paracone rib, quite flat metacone labial surface and small basal tubercles are the characters of G. capricornis (Bibi and Gulec, 2008) that are very close to G. lydekkeri. The ribs on the inner side of lower molars are less pronounced in G. lydekkeri but extremely weak in G. dorcadoides and G. pilgrimi in which the inner surfaces of the lower molars are almost flat, much like the living forms of Gazella. The posterior cleft of the p4 of G. lydekkeri is restricted to the half of the tooth towards the base of the crown, whereas in G. capricornis it is well distinguishable down towards the base. The posterior valley in p4 of G. pilgrimi is more blocked than in G. lydekkeri. The p4 of G. dorcadoides seems to resemble closely to that of G. lydekkeri. Gazella lydekkeri is different from G. capricornis in skull and horn core characters. In case of G. lydekkeri, basioccipital is comparatively narrower, horn cores are less upright, less curved and have narrower cross section as opposed to G. capricornis. Gazella lydekkeri differs from G. soriae (Alcalá and Morales, 2006) by its smaller size and greater hypsodonty but similar in the reduction of the premolar row with respect to the molars. Gazella lydekkeri shares hypsodonty and short premolar sequence as a common character with G. gaozhuangensis
(Chen, 1997) but distinct in p4 morphology and posterolabial crest of M3. 91
Figure 20. Gazella lydekkeri. 1. PUPC 83/319 left horn-core. 2. PC-GCUF 09/45 left horn-core. a) anterior view, b) medial view, c) lateral view, d) posterior view. 3. PUPC
08/136 right isolated m2: occlusal view. 4. PUPC 07/13 isolated left m3. 5. PUPC 09/76 isolated left m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
92
Discussion
The most species rich and wide spread antilopine is Gazella. Gazella and the Antilopini became widespread in the upper Miocene, probably from their possible African or
Siwalik origin (Gentry, 1994) and the fossils attributed to this taxon go back to the mid
Miocene (Gentry, 1970). Gazella became widespread in Eurasia in the late Miocene, originating probably from Homoiodorcas (Gentry, 1999). The genus Gazella was erected by Blainville (1816), who recorded the first fossil horn-cores of Gazella stehlini in the
European upper Vindobonian (Gentry, 1966). Most species of the Gazella are found as
fragmentary material, such as isolated teeth and horn cores. All the European species are
small. Pilgrim (1937, 1939) described two species of Gazella from the sediments of the
Middle Siwaliks, G. lydekkeri, and ?G. superba, on the basis of the horn core size
variation and the curvature. The holotype of G. lydekkeri is almost complete, comprising
a skull and a conjoined mandible (AMNH 19663). The studied material defines sufficient
similarity with G. lydekkeri. Pilgrim (1939) did not consider the variability owing to lack
of sufficient material. Morphological horn core variability within a species is due to the
great changes in horn core shape (Kostopoulos, 2005) and should be credited to
interaspecific variability (Fig. 21). The sample of the type locality confirms the presence
of G. lydekkeri in the Dhok Pathan Formation.
G. lydekkeri shows horn core morphology with the Eurasian G. capricornis (Fig. 21). The
horn core exhibits the characteristic features of G. capricornis (Pilgrim, 1937; Senyurek,
1954; Pilgrim and Hopwood, 1928). Recently, Kostopoulos (2005) observed the same
horn core pattern as in G. lydekkeri. Earlier authors (Gentry, 1970, 1971; Bibi and Gulec,
2008) already proposed affinity of the Siwalik G. lydekkeri with Pontian G. capricornis. 93
G. gaudryi G. paotehensis Gazella Horn-core 26 24 G. baturra G. lydekkeri Reffered 22 20 18 G. sp. G. cf. capricornis 16 14
DT 12 10 G. lydekkeri Studied 8 6 4 2 0 0 2 4 6 8 1012141618202224262830 DAP
m2 m3 12 12
10 10 8 8 6 6 Width Width 4 4 2 2 0 024681012141618 0 02468101214161820222426 Le ngth Length
Figure 21. Scatter diagram showing dental proportions of Gazella. Referred data are taken from Pilgrim (1937, 1939), Akhtar (1992) and Khan (2008), Bibi and Gulec (2008) and Kostopoulos (2005, 2009).
94
Subfamily HIPPOTRAGINAE Gray, 1821
Tribe REDUNCINI Gray, 1821
Genus KOBUS A. Smith, 1840
Type Species: Kobus ellipsiprymnus (Ogilby, 1833).
Generic Diagnosis (Pilgrim, 1939; Gentry, 1980): Reduncine of moderate to small size; with large, relatively divergent, and backwardly tilted, slightly curved horn-cores, having a slight clockwise twist, with an oval, laterally compressed cross section, insertions at a low inclination; face somewhat bent down on the cranial axis; supraorbital pits rather large and deep with pointed posterior end.
Stratigraphic Range: Middle Siwaliks.
Geographic Distribution: Late Miocene to Pliocene of the Subcontinent and Eurasia
(Pilgrim, 1939, Gentry, 1980; Thomas, 1980).
Kobus porrecticornis (Lyddeker, 1878)
Type Specimen: Frontlet with the horn-cores: GSI B229.
Emended Diagnosis (modified from 1939): A faint keel like structure is present proximally. The horn-cores are curved moderately backwards having a slight clockwise twist. Horn-cores are rather slender and decrease gradually in diameter towards the tip; their basal cross section is subovate. The surface of the horn-cores is furnished with longitudinal furrows. The presence of longitudinal furrows is variable. The lateral surface of the horn-cores is slightly flat and the medial one is convex. 95
Studied Material: Horn-cores: PUPC 87/124 A+B, left and right horn-cores, proximally
preserved fragments, perhaps same individual. PUPC 02/126 and PUPC 87/180, partial
left horn-cores.
Description
The recovered material of Kobus porrecticornis comprises only horn-cores (Fig. 22).
Horn-cores
The surface of the horn-cores bears continuous longitudinal ribs. Most of these are
remarkably strong. The cross section is ovate to subovate. The horn-cores are slightly
twisted in a clockwise direction. The horn-cores are curved backwardly and weakly
mediolaterly compressed. The mediolateral compression persists towards the apexes.
Their medial face is convex at the base while the lateral one is slightly flat. The horn- cores become tapers towards the tips (Fig. 22(1-4)). The longitudinal furrows are strongly grooved anteriorly in PUPC 87/124. The dimensions of the horn-cores at the base are:
PUPC 02/126 – DAP 32.4 mm, DT 27.6 mm; PUPC 87/180 – DAP 36 mm, DT 29 mm.
The mean index at the base DT × 100/DAP for PUPC 87/124A is 72%, PUPC 02/126 is
85% and PUPC 87/124B is 80.5%.
Comparison
The morphometric characters of the sample allot the medium size antelope. The horn- core’s slight clockwise twist excludes the sample to Gazella (Fig. 22(1-4)). Nevertheless, the torsion in Gazella may be observed in the horn tip of some species of Gazella but this does not extend to the horn-cores (Pilgrim, 1939). The material is distinguished from G. 96
rodleri in having clockwise twist because G. rodleri have anticlockwise twist (Pilgrim,
1939). The described horn-cores are clearly differentiated from those Antilope,
Helicotragus and Lithocranius in absence of strong torsion (Fig. 22(1-4)).
The overall direction, compression, slight twist and sculpture of the specimens are pretty fit for the species. The basal proportions of the studied horn-cores indicate variation in compression but this may be due to allometric growth of the horns (e.g. see description).
The described sample is being closer to reduncines Kobus in regard to the degree of
torsion, backward tilt, divergence and sculpture of the horn-cores. The studied specimens
indicate have a faint keel than that of the early reduncines K. subdolus Gentry, 1980 and
K. sp. 2 at Langebaanweg, the Redunca aff. darti of Lehmann, and K. porrecticornis
(Lydekker, 1878) of the Siwaliks and a character that could be explained by the earlier
stage of the antelope. However, the studied specimens differ from the common African
reduncine by being more transversely compressed, having less of a flattened lateral
surface, a less pronounced posterolateral angle and perhaps by having less strong
longitudinal grooving posteriorly. The horn-cores are like the Siwaliks K. porrecticornis
(Lydekker, 1878), which is the most probably from the upper part of the Dhok Pathan
Formation. Thomas (1980) noted that K. porrecticornis (Lydekker, 1878) appears
between sandstones 6 and 7 at the base of the Kundrali unit in the type zone of the Dhok
Pathan. The findings of the studied horn-cores in the Dhok Pathan type locality confirm
its stratigraphic occurrence in the upper part of the Middle Siwaliks.
Discussion
The common reduncine horn-cores vary quite a lot in size. They show little transverse
compression, the lateral surface more flattened than the medial one, their widest 97
mediolateral diameter lying rather anteriorly, a strong posterolateral angle, deep
longitudinal grooving posteriorly, transverse ridges on their anterior surfaces, insertions above the backs of the orbits, insertions at a low angle in side view, much divergence,
little backward curvature in their proximal parts, slight upward curvature distally, limited
sinus development in the vicinity of the supraorbital pit but no sinuses in the pedicels,
and deep postcornual fossae. The frontals are quite deep at the suture between the horn-
core bases, and the dorsal part of the orbital rim projects quite strongly (Gentry, 1997).
The braincase is moderately long, the cranial roof is steeply inclined but not curved
downwards in its posterior part, the mastoids are moderately sized and face partly
laterally, the ventral rim or edge just below the mastoid is weak and there is no dorsal rim
(Gentry, 1985), the nuchal crest is quite strong and forms the effective boundary of the
occipital surface, the occipital edge is not rounded in the form of a semicircle, a median
vertical ridge was present on the occipital, the temporal lines approach fairly closely, and
the auditory bulla was probably large.
The main tooth characters of living reduncines are: moderate hypsodonty; teeth rather
small in relation to skull and mandible size; upper and lower molars with
entostyle/ectostylid; lingual lobes of upper molars and labial lobes of lowers constricted;
upper molars with strong styles; localized ribs between the styles; central fossettes of
upper molars with indentations (spurs) of their outlines and tending in later wear to have
long, labially directed, transverse arms; lower molars with goat folds and front and rear
outbowings of lingual walls; upper and lower P2s small; lower premolars with an
appearance of anteroposterior compression; P4s with a strongly projecting hypoconid and
often a deep and narrow labial valley in front of it; p4s usually without paraconid- 98
metaconid fusion to form a complete lingual wall (Gentry, 1997). In general earlier
reduncine teeth are less distinctive than later ones in having lower crowned teeth,
entostyle/ectostylid smaller or even absent, less constricted lingual lobes of upper molars and labial lobes of lower molars, ribs on upper molars less localized, central fossettes of
upper molars less complicated, smaller goat folds on lower molars, lingual walls of lower
molars less strongly outbowed, longer premolar rows and the entostylid more distinct
from the entoconid on the back of p4. The Siwaliks K. porrecticornis indicates the earlier
features of reduncines and is considered more primitive than those of African reduncines
(Gentry, 1997).
The Siwalik Kobus holotype GSI B229 named Dorcadoxa porrecticornis by Pilgrim in
1939 and placed in the Antilopini instead of Reduncini. Pilgrim (1939) had followed
Lydekker and suggested a relationship of the holotype with Aepyceros considering it the
first representative of African Aepyceros. He suggested that Dorcadoxa was the ancestral
form of the living African Aepyceros and it may have been one of the early members of
antelope, having migrated to Pakistan in the Miocene (Pilgrim, 1939). Gentry (1970)
suggested a reduncine affinity. Thomas (1980) and Gentry (1980) treated the species as
belonging to early member to Kobus. Horn-cores of K. subdolus, an earlier member of
African Kobus may show an evolutionary advance over K. porrecticornis in that they are
more fully reduncine in characters such as a less compressed cross section, a more
flattened lateral surface, a more posterolateral keel, and a diminished backward curvature
basally. The studied sample reflects the primitive features of reduncine and consequently,
K. porrecticornis is the earliest member of Reduncini recovered from the sediments of
Dhok Pathan type locality of the Dhok Pathan Formation. 99
Figure 22. Kobus porrecticornis. 1. PUPC 87/124A left horn-core. 2. PUPC 87/124B right horn-core. 3. PUPC 02/126 left horn-core. 4. PUPC 87/180 left horn-core. a) anterior view, b) medial view, c) lateral view, d) posterior view. Scale bar equals 30 mm total. 100
cf. Kobikeryx sp. Pilgrim, 1939
Studied Material: Horn-cores: PUPC 99/49, distally preserved fragments, side indeterminate.
Description
Horn-core
It is not known from how far above the base it comes but it looks to break from the base
(Fig. 23). The length of the fragment is 63 mm. The horn-core appears to be less compressed medio-laterally. The slight degree of compression probably increases above the base. There is no flattening of the surfaces of the horn-core and no keels. The horn- core is tilted back and the anterior border is slightly convex. The cross section is elongate oval without keels and the surface is ornamented with irregular longitudinal furrows. A prominent longitudinal furrow presents posteriorly (Fig. 23).
The dimensions of the horn-core at the base are: PUPC 99/49 – DAP 25 mm, DT 20 mm giving the mean index DT × 100/DAP is 80%. The mean index at the base of already recovered specimens from the Dhok Pathan Formation (Pilgrim, 1939): for GSI B827 is
70%, for GSI B826 is 75.5% and for BMNH 15482 is 76.2%.
Comparison and Discussion
The reduncines show little transverse compression, the lateral surface more flattened than the medial one, their widest mediolateral diameter lying rather anteriorly, a strong posterolateral angle, deep longitudinal grooving posteriorly, little backward curvature in their proximal parts (Gentry, 1997). The studied sample reflects the characteristics of 101 reduncines and metrically accords well with the Siwalik Kobikeryx recovered from the
Dhok Pathan Formation. However the material is not enough for the specific determination.
Figure 23. cf. Kobikeryx sp. 1. PUPC 99/49 horn core, side indeterminate: a) anterior view, b) posterior view. Scale bar equals 30 mm total.
102
Subfamily CAPRINAE Gray, 1821
Tribe CAPRINI Gray, 1821
Genus CAPROTRAGOIDES Thenius, 1979
Type Species: Caprotragoides potwaricus (Pilgrim, 1939).
Generic Diagnosis (Pilgrim, 1939; Anzana and Morales, 1994): Small size, skull
relatively wide at the orbits and narrow at the brain case; face bent down on the parietal
surface at an angle of no more than 30°; horn-cores rather small, far apart, relatively
rather upright, inclined to the face at an angle of about 150° and to the parietal surface at
an angle of about 60°, somewhat oblique to the skull axis, slightly curved to the rear,
cross section elliptical, laterally compressed, without keels, slightly more convex on the
medial than on the lateral side.
Stratigraphic Range: Lower and Middle Siwaliks.
Geographic Distribution: Middle Miocene to Pliocene of the Subcontinent and Eurasia
(Pilgrim, 1939; Thomas, 1984; Kohler, 1987; Gentry, 1990).
Caprotragoides cf. potwaricus (Pilgrim, 1939)
Type Specimen: Fragmentary skull with the partial horn-cores: GSI B630.
Emended Abbreviated Diagnosis (modified from 1939): Surfaces marked by very deep longitudinal grooving, particularly on the medial and posterior aspects; cross-section ovate, mediolaterally compressed with flattened lateral face; gently curved backwards; slight torsion (clockwise on right side), approach to a keel posteriorly and upright insertion. 103
Studied Material: Horn-core: PC-GCUF 09/44, right side.
Description
Horn-core
PC-GCUF 09/44 is the single most complete specimen, including a complete right horn core (Fig. 24). The horn-core is slightly curved backwards and tapers rapidly towards the
tip. The cross section is ovate and slightly more convex on the medial than on the lateral
side. The surface is covered with longitudinal grooves; some are comparatively deep
medially and posteriorly (Fig. 24). Two deep longitudinal furrows, starting above the
base, runs in parallel and become deep towards the tip of the horn-core approach to a keel
posteriorly. The horn-core has slight twist along the grooves. The length of the studied
horn-core is 189 mm. The transverse diameter at the base is 28 mm and antero-posterior
diameter at the base is 35 mm having mean index DT × 100/DAP is 80% at the base. The
transverse diameter of the holotype (Pilgrim, 1939) at the base is 31.5 mm and the antero-
posterior diameter at the base is 41 mm having mean index DT × 100/DAP is 77% at the
base.
Comparison
This is one of the complete horn-core of Caprotragoides potwaricus ever recovered from
the Siwaliks (Fig. 24). The specimen is much larger than the Siwalik gazelle. This
specimen is immediately distinguished from Gazella by horn cores that are relatively much longer, and stouter. Horns inserted above the orbit; little inclined against the parietal surface; moderately divergent; gently curved backwards; with surfaces marked 104 by very deep longitudinal grooving, particularly on the medial and posterior aspects; cross-section ovate (mediolateral compression) with slightly flattened lateral face; with high pedicels, highest anteriorly. The medial surface is convex. The specimen is distinguished from African Gentrytragus and Tethytragus by its backwards curvature, and less mediolateral compression (Gentry, 1990; Bibi and Gulec, 2008). The specimen is a very good match of C. potwaricus based on horn core which is relatively long, curved backwards, moderately divergent, less stout and marked by deep longitudinal grooves. In
Caprotragoides, the horn-cores are without keels, slightly compressed with a lateral surface a little flattened, slight backward curvature, the front of the pedicel higher than the back. Restricted sinuses are present in the frontols around the supraorbital pits and abutting the origin of horn-core pedicels. Tooth enamel is rugose, m1-2 have small or tiny ectostylid, those on m3 are smaller again. m3 never has a talonid fossette. Premolar rows are shorter than in Eotragus, otherwise the differences are almost non-existent.
African Caprotragoides have larger and more hypsodont teeth than those of C. potwaricus (Gentry, 1970; Thomas, 1981). Kubanotragus seems to occur as early in Asia as Eotragus in Europe (Chen, 1988) and has horn-cores of rounded cross section and slight or moderate torsion (clockwise on right side). However, both are without keels and have upright insertions (Gentry, 1990). Caprotragoides differed additionally from
Kubanotragus by the hypsodonty of its cheek teeth. Caprotragoides species from
Ngorora was larger and showed dental advances, such as a more complicated outline of central cavities on its molars and paraconid-metaconid fusion on p4 (Thomas, 1981). An earlier Caprotragoides from Nyakach had an approach to a keel on its horn-core and thereby suggested a relationship to Benicerus looks as if it had a more inclined cranial 105
roof than did Caprotragoides. The African Pachytragus differs from Caprotragoides in the inclined braincase roof, less upright horn-core insertions in side view, and less hypsodont teeth, and is difficult to envisage as a direct descendent of that Caprotragoides
(Gentry, 1990).
Discussion
Caprotragoides is a non boselaphine bovid, founded for Hippotraginae gen. indet. (cf.
Tragoreas) potwaricus Pilgrim, 1939, based on a frontlet from the Nagri Formation of the Siwaliks (Thomas, 1984). Anzana and Morales (1994) proposed the classification and new nomenclature for some mainly middle Miocene bovids. They ascribed C. potwaricus
(Pilgrim, 1939) type species for the genus, founded from the middle–late Miocene of
India and Pakistan. Thomas (1984) recovered C. potwaricus from the pre-Hipparion sediments of Indian Siwaliks. The holotype frontlet of C. potwaricus and some records of
C. gentri already extend into the upper Miocene. The present sample found in the Dhok
Pathan type locality which belongs to upper Miocene (e.g. see Barry et al., 2002) extends its presence in the Dhok Pathan Formation of the Middle Siwaliks. Prior to this, it is only known from the Nagri Formation of the Middle Siwaliks.
Caprotragoides (Caprini) is the earliest bovid other than Eotragus, Miotragocerus and
Homoiodorcas (Gentry, 1990; Thomas, 1984). These occur in pre-Hipparion faunas of
Asia, Africa, and Europe. Caprotragoides has a similar relationship to Protoryx and
Caprini (Kohler, 1987) as does Homoiodorcas to the Antilopini. Hence, if Boselaphini,
Antilopini and Caprini are successive branches of bovids (Gentry, 1992), the same would 106
be true of Eotragus, Homoiodorcas and Caprotragoides. Caprotragoides shares an immediate common ancestory with Homoiodorcas (Gentry, 1994).
Figure 24. Caprotragoides cf. potwaricus. 1. PC-GCUF 09/44 right horn-core: a) anterior
view, b) medial view, c) lateral view, d) posterior view. Scale bar equals 30 mm total.
107
Family CERVIDAE Goldfuss, 1820
Subfamily CERVINAE Goldfuss, 1820
Tribe CERVINI Weber, 1928
Genus CERVUS Linnaeus, 1758
Type Species: Cervus elaphus Linnaeus, 1758.
Generic Diagnosis (Colbert, 1935; Gentry et al., 1999): The cheek teeth evolved a
greater crown height, fusion of the outer cristae, selenodont as well as intensified and
higher. The styles are stronger than the stylids and the first molar is brachydont. The goat
fold and neocrista are either in a vestigial form or absent. The upper premolars show molarization by becoming more rounded and retaining only a weak incision on the lingual wall. The ectostylid is present.
Stratigraphic Range: Middle and Upper Siwaliks.
Geographic Distribution: Asia, North America and Europe (Romer, 1974; Savage and
Russel, 1983; Pitra et al., 2004).
Cervus cf. sivalensis Lydekker, 1884
Type Specimen: A fragment of right mandible with m2-3: GSI B215.
Diagnosis (Colbert, 1935; Khan, 2007): A large cervid with relatively hypsodont
molars. The skull and antlers resemble these portions in Cervus duvaucelli. A skull with
frontal concavity at the orbits, and the forward swell at the pedcels. The lacrymal vacuity
is smaller than in C. duvaucelli. The brow tines of the antler arise immediately above the burr, and form an obtuse angle with the beam. 108
Studied Material: Lower dentition: PUPC 04/18, right mandibular ramus with m2-3.
PUPC 09/73, right mandibular ramus with m2.
Description
Lower dentition
The molars are in middle wear and the length of the mandibular ramus PUPC 04/18 is 87 mm (Fig. 25(1)). The major conids with the roots show the exact morphology of the molars. The conids are preserved up to their pointed ends, which are running laterally
towards their respective cristids. The dentine is exposed over the crown surface. The
crown shows the comparatively strong accessory columns. The ectostylid is strong and
presents up to the middle of the transverse valley. The ectostylid, anteriorly, is connected
to the postprotocristid and to the praehypocristid posteriorly. The median ribs are weak
and the mesostylid seems to be absent. The metastylid and the entostylid are present on
the lingual side and the metastylid is stronger than the entostylid, which is projected
outwardly near the summit of the crown (Fig. 25(1)). The anterior and the posterior fossettes are prominent. The anterior fossette is formed by the union of the protoconid and the metaconid, while the posterior fossette is formed by the union of the hypoconid and the entoconid. The hypoconulid is broad and it is constricted posteriorly. The lateral lobes are not constricted. The anterior fossette is more crescentic and narrower than the posterior one. The enamel is thick and rugose. The rugosity is prominent on the labial side. The basal cingulum is present (Fig. 25(1a-c)). The comparative measurements are provided in table 9.
109
Table 9: Comparative measurements of the cheek teeth of the Siwalik Cervus in mm (millimeters). * The studied specimens. Referred data are taken from Colbert (1935), Ghaffar (2005) and Khan (2007). Taxa Number Nature/Position Length Width W/L ratio C. cf. sivalensis PUPC 04/18* rm2 21.0 13.0 0.61 rm3 29.0 13.0 0.44 PUPC 09/73* rm2 20.0 12.0 0.60
C. sivalensis PUPC 68/171 lm2 25.3 18.7 0.73
PUPC 04/21 lm3 19.0 11.6 0.61 PUPC 66/9 rm2 29.0 20.0 0.68
rm3 43.0 21.0 0.48 PUPC 83/286 lm2 8.00 9.50 1.18 lm3 26.0 11.5 0.44
PUPC 84/119 lm2 21.0 15.0 0.71
lm3 24.5 13.0 0.53 PUPC 87/279 rm3 31.0 18.0 0.58 PUPC 00/92 rm2 24.0 15.0 0.62 rm3 32.0 13.5 0.42
GSI B215 rm2 25.0 17.5 0.7 rm3 35.0 -- -- C. cf. triplidens PUPC 09/71* rP4 13.0 17.0 1.30 rM1 16.0 20.5 1.28 rM2 19.0 22.0 1.15 C. triplidens PUPC 04/25 rM1 20.5 21.5 1.04 rM2 21.5 23.0 1.06 rM3 21.0 20.4 0.97 GSI B204 rM2 20.0 23.7 1.18 rM3 18.0 21.0 1.16 PUPC 98/77 lM2 19.0 22.0 1.15 lM3 18.0 21.0 1.16 PUPC 98/77 rM2 19.0 22.0 1.15 rM3 18.0 21.0 1.16 110
Comparison and Discussion
The studied teeth are crescentic, show selenodonty and can be included in Ruminantia and Tylopoda (Fig. 25; Table 9). The teeth are too small to belong to Tylopoda and
Giraffidae, and too large for Tragulidae. The specimens, in addition have weak stylids.
Their morphology is very close to Bovidae but the absence of the stylids makes their inclusion in Cervidae. The Siwalik species of the family Cervidae belong to genus
Cervus and five species, C. simplicidens, C. triplidens, C. punjabiensis, C. sivalensis and
C. rewati (Arif and Shah, 1991).
The Siwalik cervids exhibit significant size variation in their dentition from species to species and C. sivalensis is the largest one (Khan, 2007). The studied teeth are hypsodont, have accessory columns and basal cingulum, and the pronounced anterior flange. These characters make their inclusion to C. sivalensis and C. rewati and exclude them to C. simplicidens, C. triplidens, and C. punjabiensis. The diagnostic features of the C. rewati close to C. sivalensis but it is a small size species and C. sivalensis has fairly big molars when compared to those of C. rewati (Arif and Raza, 1991) (Table 9). The teeth of C. sivalensis are large and quadrate in shape. The folds are open and the enamel is rugose.
The described specimens are well worn and have metrical and morphological similarities with the type specimen (GSI B215). The studied teeth are large enough to include them
C. sivalensis, nevertheless the samples are inadequate to identify up to specific level.
Lydekker (1884) introduced the name C. sivalensis, and it was studied later by Pilgrim
(1910), Brown (1926), Matthew (1929) and Colbert (1935). Lydekker had pointed out the resemblance of the skull and the antlers of the species to C. duvaucelli. He also mentioned that the brain case of C. sivalensis is large and the face is deep. Lydekker 111
(1880) referred the type specimen GSI B215 to C. triplidens, but later on he (1884) stated that this conjectural reference was incorrect, and the name C. sivalensis referred to the type specimen GSI B215.
Figure 25. Cervus cf. sivalensis. 1. PUPC 04/18 right mandibular ramus with m2-3: a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
112
Cervus cf. triplidens Lydekker, 1876
Type Specimen: a right maxilla with M2-3: GSI B204.
Diagnosis (Khan, 2007; Colbert, 1935): The Molars are hypsodont with large accessory
columns. The molars have thick enamel folds/layers; the fossettes also have a thick layer
of the enamel.
Studied Material: Upper dentition: PUPC 09/71, right maxillary ramus with P4-M2.
Description
Upper dentition
PUPC 09/71 (Fig. 26; Table 9) is in late wear and the roots are exposed completely on the
lingual side as well as on the labial side. The length of the maxillary fragment is 47 mm.
The cheek teeth are broad crowned. The anterior facet of the P4 is depressed due to the
contact of the P3. The P4 is horse shoe shaped and have a prominent fossette which is
wide and crescentic (Fig. 26). The parastyle is strong and there is a narrow furrow
between the parastyle and the paracone rib. The paracone rib is close to the parastyle than
the metastyle. The metastyle is inclined towards the molar. The P4 has a round lingual
wall and indistinct hypocone. The praeprotocrista is shorter and less curved than the
postprotocrista. The molars are in wear and the dentine is present on the crown surface
(Fig. 26). The enamel is weathered away owing to seasonal weathering and it can be
observed on the labial side of the metacone. The entostyle situated at the base of the
praehypocista can be seen in the transverse valley. The styles cannot be observed. The
rugosity is more prominent on the lingual aspect. The lingual cusps are broad and almost
U-shaped. The fossettes having a thick enamel layer are present. The posterior fossette is 113
broader than the anterior one. The fossettes are well preserved and crescentic in shape.
They are broad in the middle and narrow anteroposteriorly. The fossettes are continuous
from the anterior to the posterior border of the enamel (Fig. 26).
Figure 26. Cervus cf. triplidens. 1. PUPC 09/71 right maxillary ramus with P4-M2: a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
Comparison and Discussion
The molars are characterized by enamel folding, absence of the basal cingulum and the
deepness of the transverse valley. Cervus sivalensis has a well developed basal cingulum which is absent in the studied sample. The deep transverse valley and the folded enamel
exclude the sample from C. punjabiensis which have a shallow transverse valley and
unfolded enamel (Brown, 1926; Azzaroli, 1954; Arif et al., 1991). The specimen differs
from C. rewati and C. simplicidens in having a large size (Fig. 27). Morphometrically,
the studied maxillary ramus (Fig. 26; Table 9) is referred to Cervus cf. triplidens
(Lydekker, 1876), due to slight traces of cingulum, strong internal column and folded
enamel on the outer surface of teeth. 114
Cervus triplidens was discovered by Lydekker (1876) and its presence was justified by
Pilgrim (1910), Brown (1926) and Matthew (1929). The holotype comprises of the right
M2-3 and the 2nd molar is slightly worn, while the 3rd is merely a germ, as yet untouched
by wear. Thus, with the wear, the length and width ratio of the crown surface must be
changed. Brown (1926) mentioned the presence of C. triplidens in the Upper Siwaliks,
while Pilgrim (1910) mentioned its presence in the Middle Siwaliks. Lydekker (1876) did
not designate any specified stratigraphic range/locality for the holotype. The discovery of
the sample from the Dhok Pathan type locality confirms its presence in the upper part of
the Middle Siwaliks.
Lydekker (1880) and Pilgrim (1913) mentioned the Upper Siwaliks as the stratigraphic
level for the Siwalik cervids. More recently, the cervids were collected from the Dhok
Pathan Formation of the Middle Siwaliks (e.g. see Ghaffar et al., 2004; Ghaffar, 2005;
Khan, 2007) and now it is also collected from the Dhok Pathan type locality in this thesis, which evidently prove that the cervids had already entered in the Siwaliks prior to 5 Ma
(see Barry et al., 2002 for the age of the Dhok Pathan). However, a perfect material is required for the confirmation.
115
C. cf. sivalensis C.sivalensis C. triplidens C. cf. triplidens
M1 M2 25 25 20 20 15 15 Width 10 Width 10
5 5
0 0 0 5 10 15 20 25 0 5 10 15 20 25 Length Length
m2 m3 25 25
20 20 15 15 Width Width 10 10
5 5 0 0 0 5 10 15 20 25 30 35 0 1020304050 Length Length
Figure 27. Scatter diagram showing dental proportions of the Siwalik cervids. Referred data are taken from Colbert (1935), Ghaffar (2005) and Khan (2007).
116
Family Tragulidae Milne-Edwards, 1864
Genus DORCATHERIUM Kaup and Scholl, 1834
Type Species: Dorcatherium naui Kaup and Scholl, 1834.
Generic Diagnosis (Whitworth, 1958; Pickford, 2002): Tragulids of varied size.
Median metacarpals distinct; median metatarsals coalesced, but retaining separate medullae. Navicular, cuboid and external cuneiform of tarsus usually fused, but external cuneiform occasionally separate; internal cuneiform always distinct. Dental formula normally 0133/3143, p1 sometimes absent. Molar teeth as in recent tragulids, but more bunoid in larger species of Dorcatherium. Median, external, accessory stylid may be present in lower molars, more often absent. Premolars usually longer, relative to breadth, than in recent tragulids. Upper canine of male, large, recurved, trenchant.
Generic Diagnosis (Farooq et al., 2008; Rossner, 2010): The teeth are bunoselenodont to selenodont with more or less strong cingula and cingulidae and mostly strong styli and stylidae at the molars. The upper molars increase in size from M1 to M3. The lower molars show a special crest complex called the ‘Dorcatherium-fold’. It is formed by the bifurcation of the posterior slopes of the protoconid and the metaconid resulting in a ‘Σ’ shape. The premolars are comparatively long and consist mainly of the labial coni/conidae and crests. Lingual crown elements are underrepresented. At the p4 the entoconid fuses with the postprotocristid. The p3 has only a short lingual entocristid originating at the hypoconid. An exception is the P4, which is shorter and does not have an anteroposterior longish shape. The cheek teeth are high crowned. The upper molars 117
bear strongly developed labial styles. The lower molars are characterized, either by well- developed ectostylid or by a vestigial ectostylid.
Stratigraphic Range: Lower and Middle Siwaliks.
Geographic Distribution: Lower to Middle Siwaliks (Middle Miocene to early
Pliocene) (Farooq et al., 2007a), MN4 (late Early Miocene, early Karpatian, middle
Orleanian) (Sach and Heizmann, 2001) to MN13 (latest Late Miocene, late Pontian or early Dacian, late Turolian) (Arambourg and Piveteau, 1929), eventually Pliocene
(Bakalov and Nikolov, 1962), in Europe, Middle Miocene (Qiu and Gu, 1991) to Early
Pliocene (Lydekker, 1876) in Asia, late Early Miocene (Pickford, 2001) to Early Pliocene
(Pickford et al., 2004) in Africa.
Dorcatherium majus Lydekker, 1876
Type Specimen: Two upper molars: GSI B197.
Diagnosis (Farooq et al., 2008): Dorcatherium majus is a species larger than
Dorcatherium minus and is equal in size to Dorcabune anthracotherioides. It is characterized by strong parastyle and mesostyle, well-developed cingulum in upper molars and stoutly developed ectostylid.
Studied Material: Upper dentition: PUPC 03/14, isolated left M2. Lower dentition:
PUPC 01/21, isolated right m1. PUPC 01/26, left mandibular ramus with m2 and posterior portion of m1. PUPC 07/09, isolated left m2.
118
Description
Upper dentition
The molar is almost quadrate in shape (Table 10) and in an early wear (Fig. 28(1)). The
basal shelf like cingulum surrounds the lingual part of the molar and it is moderately
developed at the antero-posterior face of the molar. It is present at the entrance of the
transverse valley between the protocone and the hypocone and it is entirely absent around
the labial cones. The entostyle is absent (Fig. 28(1)). The enamel is wrinkled and shows a
uniform thickness all over the crown of the molar. The anterior and the posterior cavities
are wide and deep. The principal cusps are inclined towards the median longitudinal line
of the molar and this inclination is more conspicuous in the lingual cusps than those of
the labial ones. The protocone is more worn than the other ones. It exhibits semi-
crescentic shape, as the praeprotocrista is longer than the postprotocrista. The innermost
lingual part of the protocone is not pointed. The paracone is relatively smaller in antero-
posterior length than the metacone. The antero-buccal cusp has a strong anterior groove
descending from its apex to the base of the crown, which separates the anterior style from
the main cusp. The buccal notch is V-shaped. The parastyle is very well developed and
connected to the base of the anterior median rib. The anterior median rib is more
prominent than the posterior one. The mesostyle is very well developed and it is in the
form of an isolated pillar. The metastyle is almost equal to that of parastyle in height and
its basal part is broken. The hypocone is crescentic in shape. The posthypocrista is thin
than the praehypocrista. The posthypocrista enters the metaconus leaving the metastyle
outside the fossette and making the shape asymmetrical. Labial styles and columns are
strong (Fig. 28(1)). 119
Lower dentition
The Dorcatherium-fold is present on the distal surface of the antero-lingul conids in the
lower molars (28(2-4)). The antero-lingual cusp reaches forwards to join a small
accessory cusplet in the midline, which gives a pointed contour to the front of the crown.
On its distal surface there is a low crest that descends obliquely towards the midline
where it joins the anterior crest of the disto-buccal cusp. The metaconid and the
entoconid are higher than those of the protoconid and the hypoconid. The enamel is fairly
thick and uniform in thickness. The cingulid is present on the posterior as well as on the
anterior side. It can be observed around the hypoconid. The cingulid is weakly developed on the labial side especially at the entrance of the transverse valley along the praehypocristid. A small ectostylid is present at the entrance of the transverse valley towards the labial side, and it is closely associated with the postprotocristid. The central cavities are narrow. The protoconid is anteroposteriorly longer than the metaconid. It is crescentic in shape with the praeprotocristid and the 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 metaconid is pointed at the middle with the anterior and the posterior sloping cristids. The postmetacristid is also bifurcated in PUPC
01/21 (Fig. 28(2)) and PUPC 07/09 (Fig. 28(2)) and is slightly more worn than the praemetacristid. The praemetacristid has produced a narrow and elongated dentinal islet.
The entoconid is well developed but it is slightly damaged at the posterior side. The hypoconid is more crescentic than the protoconid. The comparative measurements are provided in table 10.
120
Table 10: Comparative measurements of the cheek teeth of D. majus and D. minus in mm
(millimeters). *The studied specimens. Referred data are taken from Colbert (1935) and Farooq et al. (2007a, 2008).
Taxa Number Nature/Position Length Width W/L ratio D. majus PUPC 03/14* lM2 19.0 19.0 1.00 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 16.4 1.04 PUPC 85/15 M2 19.0 20.0 1.05 PUPC 85/21 M2 18.0 22.0 1.22 PUPC 87/328 M2 17.7 19.0 1.07 AMNH 19302 M2 18.5 21.5 1.16 GSI B198 M2 19.6 19.6 1.00 PUPC 01/21* rm1 17.5 9.00 0.51 PUPC 01/26* lm2 17.0 9.00 0.52 PUPC 07/09* lm2 18.0 12.0 0.66 PUPC 86/02 m1 14.3 9.00 0.62 PUPC 86/05 m1 13.0 9.30 0.71 AMNH 19524 m1 13.5 9.00 0.66 GSI B593 m1 15.7 9.50 0.60 PUPC 63/243 m2 17.0 10.1 0.59 PUPC 84/115 m2 16.0 12.0 0.75 PUPC 86/02 m2 15.6 9.80 0.62 PUPC 86/05 m2 15.0 11.1 0.74 PUPC 86/152 m2 16.2 12.0 0.74 PUPC 98/61 m2 17.0 10.5 0.61 AMNH 19520 m2 17.0 10.5 0.61 AMNH 19524 m2 16.0 11.0 0.68 GSI B593 m2 17.5 10.0 0.57 D. minus PUPC 68/41 M2 11.0 13.0 1.18 PUPC 68/355 M2 10.5 11.8 1.12 121
Table 10 (Continued) PUPC 86/81 M2 10.0 12.2 1.22 PUPC 95/01 M2 10.0 11.0 1.10 PUPC 02/01 M2 10.5 11.6 1.10 AMNH 29856 M2 11.3 12.0 1.06 GSI B195 M2 11.0 12.0 1.09 PUPC 68/312 m1 9.10 5.30 0.58 PUPC 68/313 m1 8.90 5.60 0.62 PUPC 02/158 m1 10.6 6.70 0.63 GSI B594 m1 12.5 7.50 0.60 PUPC 68/294 m2 11.0 6.40 0.58 PUPC 68/311 m2 10.0 6.60 0.66 PUPC 68/312 m2 10.0 6.20 0.62 PUPC 68/313 m2 10.2 6.70 0.65 PUPC 85/59 m2 9.50 7.00 0.73 PUPC 02/158 m2 12.7 8.20 0.64 AMNH 19365 m2 13.0 12.0 0.92
Comparison and Discussion
Morphologically, the specimens show typically tragulid features. The bunoselenodonty,
isolated strong styles, basal cingulum, prominent anterior median ribs and the presence of
Dorcatherium-fold full corroborate the tragulid status of the Dhok Pathan type locality
fossils (Fig. 28). The upper molars of Dorcatherium majus are characterized by subhypsodont and selenodont teeth with isolated pillar like parastyle. The metastyle is always found weakly formed (Lydekker 1876; Pilgrim, 1915; Colbert, 1935; Farooq et
al., 2007a; Rossner, 2010). The lower molars of D. majus are high comparatively and
narrow crowned. They have ectostylids between the protoconids and the hypoconids. The
metaconid is recognized as the highest one among all the conids. Stylids are weakly or 122 improperly developed. Their anterior median ribs are well pronounced. All these characteristics exhibited by the present collection of lower dentition indicate very clearly that it belongs to the species D. majus (Colbert, 1935; Farooq et al., 2008). The material assigned to this species is morphometrically (Fig. 28; Table 10) similar to the specimens from the Siwaliks described by Colbert (1935) and Farooq et al. (2007a, 2008).
The first undoubted fossil tragulid is the early Miocene genus Dorcatherium from Africa and Eurasia. The extinct genus Dorcatherium is known with many species from Asia
(Lydekker, 1876; Matthew, 1929; Farooq et al., 2007a, 2008), Europe (Rossner, 2010) and Africa (Pickford, 2001, 2002; Pickford et al., 2004). The numerous species referred to Dorcatherium mainly differ by their size (West 1980). Four Pakistani species of the genus Dorcatherium are D. minimus, D. minus, D. nagrii, and D. majus. D. minimus, the smallest species, known only by isolated upper third molar from the Chinji Formation of the Lower Siwaliks, Pakistan and D. nagrii from the Nagri Formation of the Siwaliks are rare found and probable invalid species. Dorcatherium majus and D. minus appears consistently in the Lower and the Middle Siwaliks and metrically, these are clearly distinguished to each other in size (Table 10; Fig. 29) (Farooq, 2006, Farooq et al.,
2007a, 2008). 123
Figure 28. Dorcatherium majus. 1. PUPC 03/14 isolated left M2. 2. PUPC 01/21 isolated right m1. 3. PUPC 07/09 isolated left m2. 4. PUPC 01/26 left mandibular ramus with m2 and posterior portion of m1. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
124
D. majus D. minus
M2 m1 24 10 22 20 8 18 16 14 6 12
Width Width 10 4 8 6 4 2 2 0 0 0 2 4 6 8 10 12 14 16 18 20 0 2 4 6 8 101214161820 Length Length
m2 14 12 10 8
Width 6
4 2 0 0 2 4 6 8 101214161820 Length
Figure 29. Scatter diagram showing dental proportions of the Siwalik Dorcatherium.
Referred data are taken from Colbert (1935) and Farooq et al. (2007a, 2008).
125
Genus DORCABUNE Pilgrim, 1910
Type Species: Dorcabune anthracotherioides Pilgrim, 1910.
Generic Diagnosis (Pilgrim, 1910, 1915; Colbert, 1935; Farooq et al., 2007b): Very
large tragulids having bunodont teeth. Isolated parastyle and mesostyle, prominent
cingulum and enamel rugosity are the diagnostic characteristics of the upper molars,
whereas lower molars are characterized by their broadness, a wide talonid in the third molar, and a pyramidal protoconid with two posteriorly directed folds. The upper molars of Dorcabune anthracotherioides are characterized by their brachydonty and bunodonty.
The inner cusps of upper molars are truly selenodont, whereas the outer ones are quite bunodont and absolutely conical in their general appearance. The median rib on the labial
face of the paracone and metacone is so broad and prominent that it occupies almost all
the space between the styles. This feature is very much pronounced in the paracone, the
labial surface of which is in fact entirely rib. The parastyle and mesostyle are strong,
massive and isolated, whereas the metastyle is very weakly developed. With wear, the
mesostyle clearly displays its closer association with the metacone instead of fusing
equally to both paracone and metacone. The protocone, instead of being a simple crescent, is more pyramidal in shape and displays three equally strong folds, one
proceeding forwards and outwards, the second backwards and a third backwards with a
tendency sometimes inwards and sometimes outwards. A strong cingulum runs antero-
posteriorly, but is very much pronounced round the protocone. It often rises into a small
tubercle at the entrance of the transverse valley between the protocone and hypocone.
The enamel is heavy and has moderately fine rugosity (Pilgrim, 1915; Colbert, 1935).
The lower molars are also characterized by well pronounced brachydonty and bunodonty. 126
The anterior arm of protoconid terminates on a broad shelf almost parallel to the anterior
margin of the tooth. Entoconid is conical, producing out anteriorly a short process in the
direction of the mid line between the two anterior cusps. The hypoconid is crescentic; its
anterior arm touches to the external process of the protoconid, while its posterior arm
runs inward and completely encircles the posterior base of the entoconid.
Stratigraphic Range: Lower and Middle Siwaliks.
Geographic Distribution: The genus Dorcabune is known from the Chinji Formation of
the Lower Siwaliks, the Nagri and the Dhok Pathan formations of the Middle Siwaliks
and the Lower Manchar of Bhagothoro, Pakistan (Pilgrim, 1910, 1915; Colbert, 1935;
Farooq et al., 2007b). This genus is also known from China (Han De-Fen, 1974).
Dorcabune anthracotherioides Pilgrim, 1910
Type Specimen: A maxilla with molars: GSI B580.
Diagnosis (Pilgrim, 1910, 1915; Colbert, 1935; Farooq et al., 2007b): Dorcabune
anthracotherioides is larger than Dorcabune hyaemoschoides and almost equal to that of
Dorcatherium crassum (Pilgrim, 1915; Colbert, 1935). The lower margin of the ramus is deep in Dorcabune anthracotherioides. The mandible bears a fairly deep groove starting
beneath the p4 and propagating towards the posterior side behind the teeth. This groove
exists also in Dorcatherium majus and Dorcatherium minus and in Dorcabune latidens
but is absent in Dorcabune nagrii (Pilgrim, 1915). The upper molars of Dorcabune
anthracotherioides are very similar to that of Dorcabune hyaemoschoides and differ only
by the possession of prominent parastyle. The lower fourth premolar is slightly shorter in 127 length than the lower third premolar. The p4 is broad and consisting of three lobes, of which middle one is the highest and longest, whereas first and the last lobes are equal in length, though the third lobe is higher in unworn condition. Third lobe is massive and crescent-shaped facing towards the inner and the anterior sides. The posterior arm of the crescent is running out to a level with the internal margin of the tooth. A small notch separates this arm from a long wing which runs backward from the summit of the principal cusp and forms the inner wall of the tooth. This wing is separated by a deep elongated cavity from the crest, which connects the principal cusp to the hinder lobe
(Pilgrim, 1915). Dorcabune anthracotherioides differentiates Dorcabune latidens by characterizing a less deep mandible bearing moderately broader molars and possessing much smaller size (Pilgrim, 1915).
Studied Material: Upper dentition: PUPC 99/338, isolated right M2. Lower dentition:
PUPC 04/01, right mandibular ramus with m3.
Description
Upper dentition
The molar is in an early wear, squarish, bunodont and broad crowned (Fig. 30 (1); Table
11). The enamel is extensively wrinkled and thick. The cingulum is moderately developed at the antero-posterior face of the tooth and heavily developed at the base of the protocone especially at the entrance of the transverse valley. The principal cones are inclined towards the median longitudinal line of the molar. The labial cones are relatively higher vertically than the lingual ones. The protocone is semi-crescentic in shape and displays three equally strong folds, one proceeding forwards and outwards, the second 128 backwards and a third backwards with an inwards tendency. The innermost lingual part of the protocone is pointed. The postprotocrista enters the posterior part of the anterior fossette leaving the postparacrista outside the fossette. The praeprotocrista is larger than the postprotocrista and it is continuous with the parastyle of the paracone. The hypocone is relatively smaller in antero-posterior length than the protocone (Fig. 30(1)). The praehypocrista is very small, not fused with the postprotocrista and the praemetacrista, and consequently it is free distally and mesially by deep furrows. The posthypocrista slopes downwards larger than the praehypocrista and fuses with the posterior cingulum.
The paracone is comparatively higher than the protocone. It is almost spindle-shaped, with thickness in the middle. The anterior median rib, heavy pillar like robust body siutuated labially of the paracone is separated to parastyle by a vertical groove. The metacone is smarter than the paracone. The praemetacrista is connected with the postparacrista and the postmetacrista is linked with the posthypocrista through a small ridge. It is transversely thick in the middle unlike the paracone as its median rib is much weaker than that of the paracone. The styles are moderately developed. The parastyle is well developed and it is connected to the base of the anterior median rib. The mesostyle is strongly developed and it is in the form of an isolated pillar. It seems to be more associated with the metacone than the paracone. It is thin towards its apex and very thick at the base. The metastyle is weakly developed. The central cavities are wide and very deep. Both central cavities are continuous with one another. Of these cavities, the anterior one is closed at its anterior end and the posterior one is closed at its posterior end. The posterior central cavity is also continuous with the transverse valley. The transverse valley is deep and a narrow (Fig. 30(1)). 129
Lower dentition
The third lower molar is in a good state of preservation and a bunodont (Fig. 30(2)). The
metaconid and the entoconid tips are slightly damaged. It is in an early wear. It is
hypsodont and narrow crowned. The enamel is heavy, thick and very rugose. The
cingulid is well developed anteriorly and it is absent labially, lingually and posteriorly. A
small singular tubercle is also present between the hypoconid and the talonid on the labial
side. The praeprotocristid terminates in a broad shelf, almost parallel to the anterior
margin of the tooth. The postprotocristid is bifurcated and one limb of the bifurcation is
attached to the postmetacristid while the other one is attached to the praehypocristid
producing M structure (Fig. 30(2)). The hypoconid is crescentic in shape; the
praehypocristed touches the postprotocristid, whereas the posthypocristid runs inwards
and completely encircles the posterior base of the entoconid. The metaconid is conical
and bunodont. The entoconid is conical with a short anterior process proceeding between
the two anterior conids. There is a wide vertical groove between the metaconid and the
entoconid lingually. The talonid is broad bearing a crenulated tubercle. The lingual and the labial arms of the talonid are almost equal in height.
130
Table 11: Comparative measurements of the cheek teeth of Dorcabune anthracotherioides and D. nagrii in mm (millimeters). * The studied specimens. Referred data are taken from Colbert (1935), Farooq (2006) and Farooq et al. (2007b). Taxa Number Nature/Position Length Width W/L ratio D. anthracotherioides PUPC 99/338* rM2 19.4 22.0 1.13 PUPC 87/37 M2 17.5 17.7 1.01 AMNH 19652 M2 18.0 22.5 1.25 GSI B580 M2 21.7 26.7 1.23 PUPC 04/01* rm3 29.5 15.0 0.50 PUPC 85/28 m3 26.0 13.0 0.50 AMNH 19353 m3 28.0 14.0 0.50 GSI B682 m3 30.9 16.0 0.51 D. nagrii PUPC 96/45* lM1 14.0 16.0 1.14 PUPC 68/425 M1 15.2 17.6 1.15
Figure 30. Dorcabune anthracotherioides. 1. PUPC 99/338 isolated right M2. 2. PUPC 04/01 right mandibular ramus with m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total. 131
Comparison and Discussion
The described molars exhibit buno-selenodonty pattern. The selenodonty are found in families of
Bovidae, Cervidae, Giraffidae and Camilidae and the semi-selenodonty with bunodont pattern is
found in family Tragulidae. The studied specimens reflect semi-selenodonty with bunodont
pattern and belong to family Tragulidae. There are two Siwalik genera Dorcatherium and
Dorcabune of Tragulidae (Colbert, 1935; Farooq et al., 2008). Dorcatherium is somewhat
selenodont whereas Dorcabune reflects bunodonty pattern (Fig. 28 and 30). The bunodont conical
cusp pattern of the studied samples makes its inclusion to genus Dorcabune (Fig. 30). The described upper and lower dentitions reflect morphometric characteristics of D.
anthracotherioides. The studied molars are compared with the holotype specimens as
well as the earlier described specimens (Table 11; Fig. 31) and lead to the conclusion that
the specimens are attributable to D. anthracotherioides (Colbert, 1935; Farooq et al.,
2007b).
The genus Dorcabune was founded by Pilgrim (1910) and he regarded D.
anthracotherioides as its type-species, from the Chinji Formation of the Lower Siwaliks.
Dorcabune is characterized by its very bunodont molars with thick and wrinkled enamel and the presence of the ‘double fold’ (Pilgrim 1915: 226) on the protocone of the upper molars. Pilgrim (1915) erected three more species of Dorcabune D. hyaemoschoides, D. nagrii and D. latidens from the Siwalik continental deposits. He (Pilgrim, 1910, 1915) discovered D. anthracotherioides and D. hyaemoschoides from the Chinji Formation of the Lower Siwaliks, D. nagrii from the Nagri Formation of the Middle Siwaliks and D. latidens from the Dhok Pathan Formation of the Middle Siwaliks. Colbert (1935) considered two valid species of Dorcabune D. anthracotherioides and D. nagrii in the
Siwaliks. Farooq et al. (2007b) recovered D. anthracotherioides from the Hasnot 132
outcrops of the Middle Siwaliks. The Hasnot outcrops belong to the upper portion of the
Dhok Pathan Formation (see Barry et al., 2002 for age) and now, in this study, D. anthracotherioides recovered from the Dhok Pathan type locality confirming its presence in the whole range of the Dhok Pathan Formation of the Middle Siwaliks. Based on the morphology of the lower molars of some selected genera, Moya-Sola (1988) retained
Tragulus, Hyemoschus, Dorcatherium, and, possibly, Dorcabune and Yunnanotherium in the Tragulina and excluded Archaeomeryx, leptomerycids, and hypertragulids from the
Ruminantia. Nevertheless, this point of view conflicts with the opinion of most other researchers (Gentry, 1978). According to Barry et al. (2002) the range of Dorcabune in the Siwaliks is limited to 14.0-8.4 Ma. More recently, the recovered specimens of D. anthracotherioides from the Hasnot near the Bhandar Bed (Farooq et al., 2007b) and the findings of the species in the Dhok Pathan type locality in this thesis extend its range from 14.0-5 Ma (Pilbeam et al., 1977; Johnson, N. et al., 1982; Barry et al., 1982).
D. anthracotherioides
M2 m3 30
25
20
15 Width 10
5
0 0 10203040 Length
Figure 31. Scatter diagram showing dental proportions of the Siwalik Dorcabune.
Referred data are taken from Colbert (1935) and Farooq et al. (2007b). 133
Dorcabune cf. nagrii Pilgrim, 1910
Type Specimen: A right M3: GSI B590.
Diagnosis (Pilgrim, 1915; Colbert, 1935; Farooq, 2006): Dorcabune nagrii is smaller
than D. anthracotherioides with less developed cingula. The mandible groove beneath
the teeth is absent in D. nagrii.
Studied Material: Upper dentition: PUPC 96/45, an isolated left probably M1.
Description
Upper dentition
The molar shows selenobunodonty and brachydonty (Fig. 32). The enamel is thick and rugose. The cingulum is shelf like covering the base of the protocone. It is absent on the labial and the posterior sides. The transverse valley is deep and the tubercle like cingulum is present at its enterance. The lingual cones show selenodonty whereas the labial ones seem to be bunodont. The protocone is wide in the centre and the praeprotocrista abruptly becomes narrow and elongated, and ultimately it reveals continuation with the praeparacrista. The postprotocrista is broad and enters the anterior cavity leaving free the praehypocrista. The paracone is higher than the protocone. The praeparacrista is is attached to the parastyle and the postparacrista is continuous with the praemetacrista. The paracone is supported by the prominent anterior median rib labially. The metacone has a weak median rib. The praemetacrista is fused with the mesostyle and the postmetacrista with the metastyle (Fig. 32). The praehypocrista is continuous with the postprotocrista and the posthypocrista is free and gradually narrows down posteriorly. The anterior and 134
posterior central cavities are isolated and very much reduced in length and width. The
parastyle is very much massive and isolated. The mesostyle is also well developed while
the metastyle is weakly developed (Fig. 32).
Comparison and Discussion
The molar is selenobunodont and shows the diagnostic features of the genus Dorcabune (Farooq,
2006). There are two valid Siwalik Dorcabune species D. antheracotheroides and D. nagrii
(Colbert, 1935; Farooq, 2006). The former species is larger than the latter one (Table 11). The
studied specimen is small in size, broad crowned, bunodont, and have isolated parastyle and
mesostyle (Table 11). The median ribs are well pronounced and the enamel is extensively rugose.
These characteristics assure its inclusion to D. nagrii because all the characteristics are present in the described specimen confirming their identity of the small Siwalik Dorcabune species D nagrii. Nevertheless, the matrial is very scarce and D. cf. nagrii can be assigned for the semi- selenodont molar.
Dorcabune is known in the Miocene and the Pliocene of the Siwaliks (Metais et al.,
2001; Farooq et al., 2007b). Pilgrim (1915) suggested that Dorcabune was
morphologically close to Dorcatherium and more primitive than the latter. Dorcabune
combines a set of original characters that makes it unique among all described tragulids.
Dorcabune is characterized by its very bunodont molars with thick and wrinkled enamel
and the presence of the ‘double fold’ (Pilgrim 1915: 226) on the protocone of the upper
molars. Pilgrim (1915) also pointed out a well-developed cingulum, a premetacristid
closing the trigonid anteriorly and a well-pronounced M structure with a double fold on
the posterior side of the protoconid on the lower molars. According to Qiu & Gu (1991), 135
some of the European species of Dorcatherium might be assignable to Dorcabune.
Although Colbert (1935) considered its unquestionable tragulid affinities, Gentry (1978)
claimed that Dorcabune might be a primitive anthracothere. With the exception of the M
structure and the presence of a weak entoconidian groove, Dorcabune does not display other apomorphies with Archaeotragulus, and according to Gentry (1978) the tragulid status of Dorcabune can be questioned, however a number of collected dental specimens from the Middle Siwaliks after Pilgrim (1910, 1915) evidently prove its inclusion in
Tragulina (e.g. see Farooq et al., 2007b).
Figure 32. Dorcabune cf. nagrii. 1. PUPC 96/45 isolated left probably M1: a) occlusal
view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
136
Family GIRAFFIDAE Gray, 1821
Subfamily SIVATHERIINAE Zittel, 1893
Genus HYDASPITHERIUM Lydekker, 1876
Type Species: Hydaspitherium megacephalum Lydekker, 1876.
Abbreviated Generic Diagnosis (Matthew, 1929; Colbert, 1935; Bhatti, 2005): Teeth large, quadrate, with rugose enamel. Lower teeth broader and somewhat longer.
Stratigraphic Range: Middle Siwaliks.
Geographic Distribution: South Asia (Indian subcontinent). The occurrence of the genus is recorded from various localities of the Potwar Plateau (Middle Siwaliks) of
Pakistan i.e., Dhok Pathan, Bhimber, Hasnot and it is also recovered from the Pliocene sediments of India (Lydekker, 1876; Pilgrim, 1910; Matthew, 1929; Colbert, 1935;
Bhatti, 2005; Khan, 2007).
Hydaspitherium cf. megacephalum Lydekker, 1876
Type Specimen: A skull: GSI D150.
Emended Abbreviated Diagnosis (modified from Colbert, 1935; Khan, 2007): The parastyle of Hydaspitherium megacephalum is prominent as compared to the mesostyle and the metastyle. The interior median rib is moderately developed, whereas the posterior median rib is weakly developed. The enamel is rugose. The stylids are present and the anterior fossette is transversely compressed. H. megacephalum generally is smaller than those of H. grande and H. magnum. 137
Studied Material: Lower dentition: PUPC 09/78, left mandibular ramus with broken m1, complete m2 and partially erupted m3.
Description
Lower dentition
The material comprises a complete series of lower molars (Fig. 33; Table 12). PUPC
09/78 is a fragile and it is furnished with numerous cracks. PUPC 09/78 looks a subadult animal as the m1 in early wear, the m2 unworn and the m3 semierupted (Fig. 33).
Furthermore, the mandible (PUPC 09/78) is shallow and its depth at the m2 base is 42
mm and the length of the mandibular ramus is 131 mm. The molars are in early wear, rugose
and hypsodont. The conids are not interconnected with each other. The ectostylid is absent in the
transverse valley. The cingulum is poorly developed. The fossettes are deep and narrow. The
fossette between entoconid and hypoconid is closed at its rear end. The protoconid is V-shaped
and the metaconid is spindle-shaped. The praehypocristid is longer than the posthypocristid
and both the cristids are simple. The mesostylid is more developed than those of the metastylid and the entostylid. The anterior rib is more prominent than the posterior one. The hypoconulid is not fully erupted yet and it is lower in height than the other major conids (Fig. 33).
138
Table 12: Comparative measurements of the lower cheek teeth of the giraffids in the Dhok Pathan type locality. *The studied specimens. Referred data are taken from Matthew (1929), Colbert (1935) and Bhatti (2005). Taxa Number Nature/Position Length Width W/L ratio H. cf. megacephalum PUPC 09/78* lm1 32.0 19.5 0.60
lm2 39.0 25.0 0.64
lm3 - - -
H. megacephalum PUPC 97/17 rm2 32.5 19.0 0.58
rm3 44.0 19.0 0.43
AMNH 19669 lm2 38.0 28.0 0.73
lm3 50.0 28.5 0.57
PUPC 95/24 lm3 44.0 20.0 0.45
H. cf. grande PUPC 09/77* rm2 37.0 28.0 0.75
rm3 54.5 27.0 0.49
?H. sp. PUPC 83/267* rm2 38.7 30.6 0.79
rm3 58.0 27.5 0.47
H. magnum PUPC 67/195 rm2 50.0 34.0 0.68
rm3 68.0 32.0 0.47
Comparison
The dental giraffid remains are difficult to assign at species level since size is in practice the only distinguishing criterion (Gentry, 1997). The described molars are typical giraffid molars showing basic giraffid features like the depth of the enamel folds, the obliquity of the barrels, the rugose sculpture of the enamel (Fig. 33). The teeth are large in size and can be excluded from the genus Giraffokeryx. In addition, flat entoconid wall and 139 brachydonty present in Eurasian palaeotragines, the characters are absent in the studied sample. The studied molars are characterized by more narrowly pointed labial lobes, transverselly compressed anterior fossette, hypsodonty and out bowing of lingual ribs.
These seemingly primitive characters give the teeth more resemblance to the upper
Miocene Siwalik sevatheres. Among the large upper Miocene giraffids the genera
Hydaspitheium, Giraffa, Sivatherium, Vishnutherium, and Bramatherium are possible candidates. Giraffa differs from the studied sample in brachydont teeth, lower molar set less obliquely to axis of jaw and not elongated, and the molars with ectostylid. The material differs from Sivatherium in the more pointed labial lobes (protoconid and hypoconid), more outbowing lingual ribs and the strong mesostylid (Gentry, 1997).
Vishnutherium and Bramatherium are small in size from the studied sample (Matthew,
1929; Colbert, 1935).
In the described specimen the anterior fossette is compressed transversely, the stylids are present, the anterior median rib is prominent and the enamel sculpture is coarse (Fig. 33).
These characteristics are found in the genus Hydaspitherium (Pilgrim, 1911; Matthew,
1929; Colbert, 1935; Bhatti, 2005). So the described specimens belong to the genus
Hydaspitherium. The described lower molars are clearly smaller than those of H. grande and H. magnum (Fig. 36; Table 12). The studied material’s dimensions close to species
H. megacephalum. The described specimen and the referred specimens of H. megacephalum appear same in size and crown structure (Pilgrim, 1911; Colbert, 1935;
Bhatti, 2005). So the material belongs to the genus Hydaspitherium and more specifically to the species H. cf. megacephalum which is one of the smallest species of the genus. 140
Figure 33. Hydaspitherium cf. megacephalum. 1. PUPC 09/78 left mandibular ramus with broken m1, complete m2 and partially erupted m3: a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
Hydaspitherium cf. grande
Type Specimen: A left upper molar: GSI B155.
Emended Abbreviated Diagnosis (modified from Colbert, 1935): A large species, considerably larger than H. megacephalum, but otherwise similar to it. The mandible is relatively smart and have mandibular groove beneath the teeth. The enamel is strongly rugose. The lower molars are broader and somewhat larger.
Studied Material: Lower dentition: PUPC 09/77, right mandibular ramus with m2-3.
Description and Comparison
The length of the mandibular ramus is 175 mm and its depth at m3 is 73 mm. A portion of the ascending ramus is also preserved which extends upwards to the rear end of the 141
talonid (Fig. 34). The mandibular ramus is relatively slim and a groove is present beneath
the teeth, which runs along the horizontal ramus. The groove is not observed in the
samples assigned to H. megacephalum and H. magnum. The lower teeth resemble to H.
megacephalum in the enamel sculpture, transverselly compressed anterior fossette,
structure of the crown, obliquely to axis of jaw, pointed labial lobes and outbowing of
lingual ribs but metrically the sample is considerably larger than that of H.
megacephalum (Fig. 36) and assign to H. cf. grande (Matthew, 1929; Colbert, 1935), as
the material is insufficient for the more specifically identification (Fig. 34; Table 12).
Figure 34. Hydaspitherium cf. grande. 1. PUPC 09/77 right mandibular ramus with m2-3: a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
142
?Hydaspitherium sp.
Studied Material: Lower dentition: PUPC 83/267, right mandibular ramus with m2-3.
Description and Comparison
This is one of the difficult specimens to identify. The length of the mandibular ramus is
158 mm, its depth at the m3 is 74 mm and the mandible maximum width is 40 mm. The sample is finely rugose and somewhat stout (Fig. 35). The mandible and teeth’s dimensions are about the same as in PUPC 09/77, here assigned to H. grande (Fig. 36;
Table 12). The sample differs from H. grande in the rugosity of the enamel, the mandible stoutness and the width of the mandible which is 33 mm in PUPC 09/77 contrary to 40 mm of the studied specimem (Fig. 35). These characters give the sample more resemblance to Helladotherium and Camelus (Matthew, 1929; Colbert, 1935).
The enamel is finely rugose in Helladotherium (Pilgrim, 1911) but Matthew (1929) stated that Helladotherium never entered in the Siwaliks. The Camelidae is not known in the
Dhok Pathan Formation and in addition, Camelus lower molars are characterized by the fine rugose enamel, flat lingual walls and the retention of the anteroexternal fold. Unlike
Camelus and Helladotherium, the lower molars are about the size of H. cf. grande, labial lobes are pointed, more outbowing lingual ribs and have weak stylids. The anterior fossette is compressed transversely and the molars are obliquely to axis of jaw.
Therefore, the problematic sample (Fig. 35) could be doubtfully referred to
Hydaspitherium sp. on the basis of its teeth characters and not to Helladotherium or
Camelus. 143
Figure 35. ?Hydaspitherium sp. 1. PUPC 83/267 right mandibular ramus with m2-3: a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
Discussion
The Siwalik giraffids comprise two groups regarding to the height of the molars, brachydonts and subhypsodont to hypsodonts. The brachydont giraffids include two Siwalik genera of middle
Miocene Giraffokeryex and Giraffa. The enamel sculpture is not fine in Giraffa as well as the major cusps are not in a straight line. The cusps are straightline in Giraffokeryx and the enamel sculpture is fine. The styles, stylids and the median ribs are very weak in Giraffokeryx whereas they they are strong in Giraffa. The crown is narrow in Giraffokeryx and it is broad in Giraffa
(Pilgrim, 1910, 1911; Colbert, 1935). Giraffa priscilla has more brachydont teeth with prominent styles (especially metastyle) than those of Giraffa punjabiensis and Giraffa sivalensis. The posterior half of the molar is much reduced transversely in Giraffa sivalensis, it is less reduced in
Giraffa punjabiensis and Giraffa priscilla. Upper premolars are relatively small and narrow in
Giraffa punjabiensis (Gentry, 1997). 144
The subhypsodont to hypsodont Siwalik giraffids include the upper Miocene genera Sivatherium,
Hydaspitherium, Helladotherium, Vishnutherium, Bramatherium, Indratherium and Giraffa punjabiensis (Colbert, 1935; Bhatti, 2005). Vishnutherium and Bramatherium are relatively small genera as compare to Hydaspitherium. In Helladotherium enamel is finely rugose and Colbert
(1935) synanymized Helladotherium grande with Hydaspitherium grande. Bhatti (2005) gave his opinion about the validity of the species but according to my point of view, it is just superlative assumption as there is a lot of variation in the Siwalik giraffid dentition and need a more reliable material to resolve the issue.
H. cf. megacephalum H. megacephalum H. cf. grande ?H. sp. H. magnum
m2 m3 40 35 35 30 30 25 25 h 20 dt 20 Width
Wi 15 15 10 10 5 5 0 0 0 5 10 15 20 25 30 35 40 45 50 55 0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 Length Length
Figure 36. Scatter diagram showing dental proportions of the studied giraffids. Referred data are taken from Matthew (1929), Colbert (1935) and Bhatti (2005).
145
Family SUIDAE Gray, 1821
Subfamily SUINAE Zittel, 1893
Genus PROPOTAMOCHOERUS Pilgrim, 1925
Type Species: Propotamochoerus hysudricus (Stehlin), 1899-1900.
Generic Diagnosis (Pickford, 1988): Medium sized suidae with facial part of skull
longer than cranial part; zygomatic arches depart from maxilla at right angles opposite
M1; shallow but extensive fossae present on dorsal and ventral surfaces of prezygomatic
shelf; snout nearly square in section; posterior choanae U-shaped, open immediately
behind M3; large canine flanges reach backwards from I3 to P2; basicranium advanced;
parietal lines not widely separated, but no sugittal crest; 2 cusplets in sagital valley of P4;
no labial or lingual cingula on P4; p4 with variable ‘innenhugel’ and low to medium
hypsodont anterior cusp and cingulum; dental formula 3143/3143; upper canine almost
surrounded by cementum; molar enamel thickness moderate (2 ± 0.4 mm); Furchen
shallow; no cingula on molars, except for occasional basal pillars in ends of median
valleys; talon/talonid of M3/m3 relatively simple, no extra cusp pairs; mandible deep and
robust; symphysis reaches back to rear of p2; canines not at corners of anterior ends of symphysis; anterior and posterior cusps of premolars variable, but never reach the same height as the main cusps.
Stratigraphic Range: Middle Siwaliks.
Geographic Distribution: Propotamochoerus occurs in the upper Miocene sediments of
the Middle Siwaliks. The age range is from about 10 million to about 6 million years, a
range that is comparable to the European Korynochoerus (Pickford, 1988). The localities 146
of the Siwalik Hills that yield Propotamochoerus remains are Hasnot, Kotal Kund, Lehri,
Kundvali, Gandakas, Utran, Dhok Mila, Jabi, Nagri, Khushalghar, Khaur, Kaulial, Mari,
Ava, Bandar, Perim Island, Haritalyangar (Pilgrim, 1926b; Matthew, 1929; Colbert,
1935; Pickford, 1988).
Propotamochoerus hysudricus (Stehlin), 1899-1900
Type Specimen: Right mandibular ramus with p3-m3 in worn condition: GSI B30.
Diagnosis (Colbert, 1935; Pickford, 1988): No diastemata in upper or lower cheek tooth rows; orbits open over talon of M3; zygomatic arches leave face at midline of M1; lower premolars compressed with a long and complex third molar.
Studied Material: Lower dentition: PUPC 07/32, left mandibular ramus with p4-m3; the p3 roots are also preserved in the alveoli. PUPC 99/02, left mandibular ramus with p4- m1. PUPC 94/65, right mandibular ramus with p4-m2. PUPC 97/15, right isolated m3.
Description and Comparison
The collected samples include the lower dentitions with mandibular rami (Fig. 37; Table
13). The best preserved specimen is labeled as PUPC 07/32 (37(1)) and has a complete series of the lower molar, m1-3; the length of the molar series is 64 mm. It is broken posteriorly from where the ascending ramus hides the talonid of m3 in side view. The ascent begins well behind m3, so that this tooth is completely visible in lateral view. The length of the mandibular ramus (PUPC 07/32) is 112 mm and the depth of the mandibular ramus at m3 is 39 mm. The length of the mandibular ramus (PUPC 94/65) is 80 mm and 147
its depth at m2 is 41 mm (Fig. 37(2)). The mandibular rami have robust bodies. The
mandible of P. hysudricus reflects these above mentioned characteristics.
The fourth premolar is a molarized tooth and the central cusp has migrated lingually to
form the innenhugel and it appears subsymmetrical in lateral view. The metaconid is
weak and occurs posteriorly to the protoconid. The posterior accessory cusplet is
enlarged and is bordered by an inflated cingulum. The anterior accessory cusp and
anterior cingulum are moderately high. The first molar is a four cusped tooth with anterior, median, and posterior accessory cusplets in the midline of the crown (Fig. 37;
Table 13). The posterior accessory cusplet is prominent and centrally placed. The molar is bunodont with the usual suid layout of four main cusps arranged in two lophs. The median basal pillar is present buccally in the median valley. The second molar is the large version of the m1. The m3 differs from the m2 by the presence of talonid and wide anterior lobes. The talonid of the m3 is moderately complex having a pair of cusplets side by side. There is moderate buccal flare in the molars, but lingual flare is minimal. The lingual cusps are slightly higher than the labial ones. The lower cheek teeth enamel is moderately thick and wrinkled.
The giant suids of the Siwalik belong to the genera Tetraconodon, Sivachoerus and
Hippopotamadon. The first two genera have characteristics inflated p3-4. The cheek teeth of the genus Hippopotamodon are extremely large. The studied material is too large to include Hippoptomadon. Propotamochoerus hysudricus is accompanied throughout the
Middle Siwaliks by the species Hippopotamodon sivalense. The specimens described here contain evidence about the species Propotamochoerus hysudricus and morpho- 148
metrically close to the material of the species, described by Pickford in 1988 (Fig. 39;
Table 13).
Propotamochoerus seems to have been derived from Hyotherium which occurs at the
Chinji Formation, and it is possible that the changeover from one form to the other occurred at about 11 million years, just where the record is poor. The main advances made by Propotamochoerus over Hyotherium are the addition of sagittal cusplets in P4, increase in size, alteration in the shape of the anterior zygomata, and squaring off of the snout section (Pickford, 1988).
149
Table 13: Comparative measurements of the cheek teeth of Propotamochoerus hysudricus and ?Hippopotamodon sp. in mm (millimeters). *The studied specimens.
Referred data are taken from Colbert (1935) and Pickford (1988).
Taxa Number Nature/Position Length Width W/L ratio P. hysudricus PUPC 07/32* lp4 14.4 10.5 0.72 lm1 16.0 12.0 0.75 lm2 20.5 15.4 0.75 lm3 28.0 16.4 0.58 PUPC 99/02* lp4 14.5 11.0 0.75 lm1 15.2 11.2 0.73 PUPC 94/65* rp4 15.0 9.50 0.63 rm1 16.0 12.0 0.75 rm2 21.0 15.0 0.71 PUPC 97/15* rm3 32.5 15.5 0.47 GSP 12343 p4 16.7 9.20 0.55 GSP 5823 p4 14.4 11.3 0.78 GSP 9355 p4 16.6 11.4 0.68 GSP 12732 p4 16.0 9.40 0.58 GSP 457 p4 15.0 11.4 0.76 GSP 5230 p4 16.4 10.8 0.65 GSP 48 p4 - 10.5 - GSP 2807 p4 14.5 10.0 0.68 GSP 7017 p4 15.6 12.6 0.80 GSP 2503 p4 13.6 11.6 0.85 GSP 10998 p4 15.5 11.6 0.74 B742 p4 15.3 10.7 0.69 B39 p4 13.9 9.90 0.71 B32 p4 18.0 12.0 0.66 B715 p4 17.5 12.5 0.71 GSP 10998 m1 17.6 13.0 0.74 GSP 2807 m1 16.0 12.0 0.75 150
Table 13 (Continued) GSP 457 m1 16.8 11.5 0.68 GSP 6226 m1 16.7 12.0 0.71 GSP 7017 m1 14.6 13.0 0.89 GSP 6727 m1 16.3 11.4 0.69 GSP 4773 m1 16.6 12.5 0.75 GSP 5112 m1 18.4 11.7 0.63 GSP 7384 m1 17.5 12.4 0.70 GSP 594 m1 16.5 11.9 0.72 GSP 10082 m1 17.0 13.0 0.76 GSP 4088 m1 18.8 12.3 0.65 GSP 149 m1 16.0 10.8 0.67 GSP 9146 m1 16.0 11.0 0.68 GSP 9355 m1 17.4 12.0 0.68 GSP 12732 m1 18.0 12.0 0.66 GSI 18082 m1 16.0 12.5 0.78 B32 m1 17.0 13.0 0.76 B715 m1 16.7 13.0 0.77 B742 m1 14.5 11.0 0.75 B50 m1 16.7 11.6 0.69 GSP 12201 m2 22.5 16.5 0.73 GSP 10082 m2 22.0 17.3 0.78 GSP 6011 m2 22.3 16.3 0.73 GSP 10098 m2 22.2 16.3 0.73 GSP 2807 m2 20.6 14.3 0.69 GSP 7384 m2 22.0 15.0 0.68 GSP 7017 m2 20.7 14.7 0.71 GSP 2503 m2 21.0 17.0 0.80 GSP 4773 m2 23.5 16.7 0.71 GSP 5212 m2 24.0 18.0 0.75 GSP 9145 m2 19.3 16.0 0.82 GSP 9403 m2 23.0 17.0 0.73 151
Table 13 (Continued) GSP 9355 m2 22.7 15.5 0.68 GSP 12863 m2 - 16.5 - B742 m2 20.0 13.9 0.69 B39 m2 19.0 14.5 0.76 B32 m2 23.0 17.0 0.73 B715 m2 23.0 16.4 0.71 GSP 9145 m3 31.3 17.0 0.54 GSP 9405 m3 34.0 19.0 0.55 GSP 9403 m3 33.5 18.0 0.53 GSP 11684 m3 32.5 18.0 0.55 GSP 9355 m3 34.5 18.5 0.53 GSP 10224 m3 34.6 17.9 0.51 GSP 10225 m3 34.7 18.8 0.54 GSP 6944 m3 32.5 17.6 0.54 GSP 10231 m3 31.4 18.5 0.58 GSP 7318 m3 35.0 18.5 0.52 GSP 2807 m3 28.5 15.9 0.55 GSP 6226 m3 31.7 16.0 0.50 GSP 7017 m3 34.5 19.3 0.55 GSP 12787 m3 32.7 17.8 0.54 GSP 7348 m3 31.8 17.0 0.53 GSP2303 m3 32.5 19.3 0.59 GSP 12201 m3 34.0 17.8 0.52 GSP 10082 m3 35.5 19.0 0.53 GSP 11007 m3 31.1 17.7 0.56 GSP 2453 m3 32.0 16.7 0.52 GSP 10998 m3 33.5 18.2 0.54 B32 m3 36.0 18.0 0.50 B715 m3 35.5 19.5 0.54 GSP B742 m3 31.0 16.0 0.51 GSP B39 m3 30.0 15.0 0.50 152
Table 13 (Continued) ?Hippopotamodon sp. PUPC 00/14* lm1 27.0 19.0 0.70 PUPC 08/26* rm2 23.0 17.0 0.73 rm3 32.0 18.0 0.56 H. sivalense B714 m1 25.0 16.8 0.67 (Pickford, 1988) GSP 4673 m1 26.8 17.5 0.65 GSP 330 m1 28.6 19.4 0.67 GSP 4086 m1 28.7 19.5 0.67 K47/798 m1 28.5 19.0 0.66 B435 m1 27.5 20.5 0.74 B2 m1 26.6 18.5 0.69 B358 m1 24.0 18.0 0.75 B25 m1 29.6 20.0 0.67 B539 m1 25.0 20.0 0.80 B435 m2 39.0 26.3 0.67 B2 m2 36.0 25.0 0.69 B358 m2 30.0 22.9 0.76 B539 m2 35.5 25.0 0.70 B714 m2 31.5 21.8 0.69 B397 m2 30.0 24.0 0.80 B21 m3 57.0 29.0 0.50 B2 m3 54.5 29.6 0.54 B358 m3 53.5 26.6 0.49 B22 m3 62.5 31.0 0.49 B539 m3 56.6 27.2 0.48
153
Figure 37. Propotamochoerus hysudricus. 1. PUPC 07/32 left mandibular ramus with p4- m3. 2. PUPC 94/65 right mandibular ramus with p4-m2. 3. PUPC 99/02 left mandibular ramus with p4-m1. 4. PUPC 97/15 right isolated m3. a) occlusal view, b) lingual view, c) labial view. Scale bar equals 30 mm total.
154
Genus HIPPOPOTAMODON Lydekker, 1877
Type Species: Hippopotamodon sivalense Lydekker, 1877.
Generic Diagnosis (Pickford, 1988): Giant Suinae in which the males have large flaring canines; molar enamel relatively thin; molars relatively simple with well developed furchenplan; labial cusps in lower molars lower crowned than lingual ones; P4 with posterior accessory cusp almost as large as two main labial cusps; posterior choanae U- shaped, open immediately behind M3; p4 with prominent innenhugel and 2-3-4 cusp, anterior cingulum and accessory-1 cusp moderately high; diastema between C-P1-2 short; broad flat dorsal surface to brain case.
Stratigraphic Range: Middle Siwaliks.
Geographic Distribution: Hippopotamodon is common in the upper Miocene of the
Siwaliks. It is almost always associated with Hipparion and its range is very similar to that of Propotamochoerus hysudricus and Microstonyx (Pickford, 1988).
?Hippopotamodon sp.
Studied Material: Lower dentition: PUPC 00/14, left mandibular ramus with broken p4 and partially complete m1. PUPC 08/26, badly crushed right mandibular ramus with damaged m1-3, probably juvenile animal.
Description and Comparison
The recovered samples are badly damaged unfortunately. However, the general contour of the molars allows to study the morphological features of the specimens. The features are based on the sample, labeled as PUPC 00/14 (Fig. 38; Table 13). The premolar crown 155 is damaged but its basal part is preserved. It is somewhat large tooth (Table 13). The molars are simple with thin enamel. The median valley is wide, the furchen moderately deep and narrow. The basal pillar is present in the median valley. However, the mandibular rami are damaged but they preserve enough to throw light on their diagnostic features. The morphological features of the samples resemble with Hippopotamodon, described from the Dhok Pathan Formation of the Siwaliks by Pickford (1988). The metrical values of the sample are smaller than those of the already described material, as the material belongs to juvenile animal and furthermore, there is a trend towards increase in size with time in Hippopotamodon, specimens from the Nagri being smaller than those from the Dhok Pathan, and these smaller than those from the Hasnot area (Pickford,
1988). The material may be assigned to ?Hippopotamodon sp. based on the morphometric features (Fig. 38-39; Table 13).
Figure 38. ?Hippopotamodon sp. 1. PUPC 00/14 left mandibular ramus with broken p4 and partially complete m1: a) occlusal view, b) lingual view, c) labial view. 2. PUPC 08/26 badly crushed right mandibular ramus with damaged m1-3: occlusal view. Scale bar equals 30 mm total. 156
Discussion
In the Siwaliks lower Miocene (20 Ma to 15 Ma) siuds are very poorly known, being
represented by two specimens, one each of Libycochoerus affinis and Libycochoerus
fategadensis. Hyotherium, Listriodon and Conohyus are represented in the Siwalik
middle Miocene (15 Ma to 10.5 Ma).
Suid diversification found considerably in the Siwalik upper Miocene (10.5 Ma to 5.5
Ma). During the early part of the period (from base of the Nagri to top of the type Dhok
Pathan) the species Propotamochoerus hysudricus and Hippopotamodon sivalense are the
most abundant, with Tetraconodon magnus, Conohyus indicus and Lophochoerus nagrii
(Pickford, 1988). In sediments younger than the type Dhok Pathan, the species
Hippohyus lydekkeri, Sivahyus punjabiensis and Sivachoerus prior make their
appearance, while both P. hysudricus and H. sivalense persist beside them.
Propotamochoerus hysudricus and H. sivalense are typically found in the Siwalik upper
Miocene and their presence may therefore be taken as good evidence of an upper
Miocene age (Pickford, 1988). It is possible to state that P. hysudricus and H. sivalense
occur from at least the middle part of the Nagri Formation, upwards to the top of the
Dhok Pathan Formation, and into younger deposits where Sivahyus and Hippohyus are
found.
157
P. hysudricus ?Hippopotamodon sp. H. sivalense
m1 m2 25 30 20 25 20 15 15 Width 10 Width 10 5 5 0 0 0 5 10 15 20 25 30 35 0 1020304050 Length Length
m3 35 30 25 20 15 Width 10 5 0 0 10203040506070 Length
Figure 39. Scatter diagram showing dental proportions of the studied Dhok Pathan
Suoids. Referred data are taken from Colbert (1935) and Pickford (1988).
158
PALAEOENVIRONMENT AND PALEOBIOGEOGRAPHY OF THE
DHOK PATHAN ARTIODACTYLS
Fossil artiodactyls are common in the Miocene and Pliocene sediments of the Siwaliks.
The Dhok Pathan paleontological study area belonging to the Middle Siwaliks, located in the Potwar Plateau of northern Pakistan, has yielded fossils of late Miocene to early
Pliocene. The Artiodactyla from the Dhok Pathan type locality are documented by outstandingly rich material. On the one hand it increases knowledge of the odontological morphometric characters of the Tragoportacina, the antelopes, the tragulids, the giraffids, the cervids and the suids. On the other hand, it completes the picture of the faunal composition of the Siwalik late Miocene artiodactyls. Despite the commonness of those species in the Siwalik late Miocene, earlier descriptions were hitherto only based mostly on the earlier material of unknown origin (e.g. see Matthew, 1929; Colbert, 1935;
Pilgrim, 1939) and the findings enable to calibrate stratigraphic position of these species with the newly discovered Miotragocerus larger sp. and ?Hydaspitherium sp.
The artiodactyls are the most abundant elements in the Dhok Pathan type locality, as in all other Eurasian late Miocene sites (Pilgrim, 1939; Pickford, 1988; Haile-Selassie et al.,
2004; Bhatti, 2005; Farooq, 2006; Bibi, 2007; Khan et al., 2008b; Bibi et al., 2009; Khan,
A. M., 2010). Most of these taxa are also known from other contemporaneous Siwalik sites (Akhtar, 1992; Khan, 2007; Khan et al., 2009a). The Dhok Pathan artiodactyl fossil record shows that the boselaphine bovids were highly diversified around the late Miocene and early Pliocene, as were the antilopines. Bovids constitute 56% of the recovered artiodactyls (Fig. 40). Out of this, 29.7% are boselaphines, 10.5% are antilopines, 10.5% 159
are reduncines and 5.3% are caprines. Cervids constitute 10.5%, tragulids are 10.5%,
giraffids are 15.7% and suids are 10.5% of the identified species from the Dhok Pathan
type locality deposits (Fig. 40).
Dhok Pathan Artiodactyls
Suidae Giraffidae 9% 13%
Tragulidae Bovidae 13% 56% Cervidae 9%
Figure 40. Percentage of species represented in the Dhok Pathan artiodactyls.
Boselaphine remains were abundant during the Eurasian, African and the Siwalik late
Miocene, and are present in the Dhok Pathan type locality. Tragoportax, Selenoportax,
Pachyportax and Miotragocerus are the dominant late Miocene Siwalik boselaphines.
Species of these genera range throughout the late Miocene (10.5 Ma – 5.5 Ma), and are
found across Eurasia from China to Spain, in the Siwaliks, and to the far south of South
Africa (Pilgrim, 1939; Moya-Sola, 1983; Thomas, 1984; Bibi, 2007; Bibi et al., 2009).
The fossil boselaphines from the late Miocene Baynunah Formation (United Arab
Emirates, Gentry, 1999) suggest an overlapping of the North African, Greco-Iranian
(=Sub-Paratethyan), and Oriental (south Asian) Provinces (Bibi et al., 2009).
Tragilaphines were absent in the Dhok Pathan type locality suggesting that the niche
probably was filled by the boselaphines. The tragelaphines became more diverse during 160
the late Miocene in Africa but the boselaphines became extinct at the end of the Miocene
in Africa (Haile-Selassie et al., 2004).
Two antelope species are identified from the Dhok Pathan type locality: Prosetrpsiceros
cf. vinayaki and Gazella lydekkeri. One caprine, Caprotragoides cf. potwaricus is documented from the type locality. Reduncines are represented in the Dhok Pathan type locality by two species. Four specimens are assigned to Kobus porrecticornis and one
specimen is assigned to cf. Kobikeryx sp. Kobus porrecticornis considers to be an early reduncine and represents from the Dhok Pathan Formation of the Siwaliks (Pilgrim,
1939; Gentry, 1980). Kobus porrecticornis dominated as a prominent element of the
Dhok Pathan type locality and it was represented by a reasonable number of specimens.
Kobus aff. porrecticornis, from the Kenyan Mpesida Beds (ca. 6.5 Ma) and the Lukeino
Formation (ca. 6.0 Ma), is considered to belong to the Siwalik Kobus porrecticornis
(Thomas, 1979; Gentry, 1980; Vrba, 1995). The earliest reduncine records in Eurasia and
in the Siwaliks date to the late Miocene (Vrba, 1995; Gentry, 1997). Late Miocene links
between sub-Sahran Africa and southern Asia are also apparent in the shared presence of bovid taxa such as the reduncine Kobus porrecticornis (Vrba and Haile-Selassie, 2006; in
this thesis).
Alcelaphines are absent in the Dhok Pathan type locality. The absence of alcelaphines
may be an indication of the absence of a vast expanse of large open grasslands. The
abundance of reduncines, antilopines and boselaphines indicates a mixture of woodland
and grassland biomes. The absence of alcelaphines from the Dhok Pathan type locality
clearly reflects ecological differences. Extant alcelaphines are known to live in arid
plains, deserts, savanna, and grassland. These habitats might have not been available for 161
the alcelaphines to flourish in the Dhok Pathan deposits. A closed habitat has been
inferred for the late Miocene boselaphines (Khan et al., 2009a). Some taxa of the Dhok
Pathan deposits inhabit variously drier bush and woodlands into moisture valleys and forests. What can be inferred from this observation suggesting a closed habitat in the late
Miocene to a more open habitat in the Pliocene.
Cervids are represented by two species from the type locality: Cervus simplicidens and
Cervus triplidens. The primitive cervine was probably tropical in distribution, it certainly inhabited woodland or open country, not closed forest, and it probably lived in eastern
Eurasia or probably India (not in western Eurasia or Southern Asia) (Kahlke, 1976;
Lister, 1984, 1986; Sommer et al., 2008).
Tragulids are identified by two genera Dorcatherium and Dorcabune from the Dhok
Pathan type locality. Dorcatherium is rather indicative of swampy-paludal habitats
(Köhler, 1993; Rössner and Mörs, 2001). Dorcatherium with cervids in the artiodactyl
community from the Dhok Pathan type locality reflects an environment of mainly closed forest. A significant representation of tragulids with other artiodactyls indicates humid
conditions, accompanying open areas (Rossner, 2010). Dorcabune anthracotherioides
were collected from the Dhok Pathan type locality and according to Barry et al. (2002)
the range of Dorcabune in the Siwaliks is limited to 14.0-8.4 Ma. More recently,
Dorcabune anthracotherioides recorded from the Hasnot (Farooq et al., 2007b) near the
Bandar Bed which is located in the east-north of Hasnot and considered the youngest part
of the Dhok Pathan Formation (Pilbeam et al., 1977; Johnson, N., et al., 1982; Barry et al., 1982). 162
Giraffids represent three species of Hydaspitherium in the Dhok Pathan type locality fossil record. These species are also known from other contemporaneous sites in the
Siwaliks and elsewhere. Hydaspitherium is the most common giraffid at the Dhok Pathan and the equivalent sites dated to between the late Miocene and early Pliocene. However,
Giraffa, Bramatherium and Vishnutherium are documented from the late Miocene of the
Siwaliks and Sivatherium is documented from younger deposits (Matthew, 1929; Colbert,
1935; Khan, 2007).
Suid remains found in the Dhok Pathan type locality are assigned to two genera,
Propotamochoerus and Hippopotamodon. The deposits exhibit the presence of the two contemporaneous and sympatric genera in the late Miocene and early Pliocene. An analogous situation occurs in Europe, where Korynochoerus and Microstonyx are common (Pickford, 1988). The known range of the genera is from about 10 Ma to about 6
Ma, but older and younger specimens might be expected to be found. These are upper
Miocene species, apparently dying out by the Pliocene.
The artiodactyl assemblage from the Dhok Pathan type locality reflects a qualitative taxonomic structure well known from other communities of Eurasia and Africa in the late
Miocene and early Pliocene (Pilgrim, 1939; Pickford, 1988; Gentry, 1997; Bibi, 2007;
Vrba and Haile-Selassie, 2006). The artiodactyl composition from the Dhok Pathan type locality is peculiar in having a comparatively high species number. The combination of a large number of specimens at the Dhok Pathan locality means that the structure of the palaeocommunity is most probably not a product of taphonomical effects, but can be taken as real. 163
Besides the here described twenty three species of artiodactyls, the perissodactyl fauna from the Dhok Pathan type locality comprises rhinos, chalicotheres and hipparions
(Heissig 1972; Iqbal et al., 2009; Khan et al., 2009b; Khan, A. M., 2010). This number is unusual in modern world biomes, where the highest number of brachydonts and hypsodonts is represented. But, it corresponds to the composition fauna of the late
Miocene of Eurasia and Africa (Gentry, 1980; Haile-Selassie et al., 2004; Kostopoulos,
2006; Bibi et al., 2009), which is associated with dry and more seasonal climate in the late Miocene. This is well in accordance with recently documented open woodlands or grassy woodlands with C4 grasses. Plant communities with predominantly C3 plants were greatly diminished after 7.0 Ma, and those with predominantly C4 plants, appeared as early as 7.4 Ma. A few paleosols may have formed under water logged, grassy woodlands, but most formed under drier conditions and closed vegetation (Barry et al.,
2002). The selenodonty of the artiodactyl groups may be interpreted for fibrous foods which in most likely may have been swamp vegetation due to the depositional environment. The combined occurrence of such taxa in the locality suggests open but no grassy habitats, including some elements of fairly abrasive vegetation (Janis et al., 2002).
164
CONCLUSIONS
The Dhok Pathan mammalian fauna suggests a much higher diversity during the late Miocene of the Siwaliks (Lydekker, 1878; Matthew, 1929; Colbert, 1935; Pilgrim, 1926b, 1937, 1939; Thomas, 1984; Akhtar, 1992; Bhatti, 2005; Ghaffar, 2005; Iqbal et al., 2009; Khan, A. M., 2010). On the basis of the faunal studies, it is concluded that during the late Miocene and early Pliocene the Dhok Pathan region was considerably more humid and better vegetated than it is today (Barry et al., 2002). The artiodactyl assemblages suggest a mix of heavy cover and, to somewhat lesser extent, forested environment. The presence in the area of some more open habitats can not be ruled out. The artiodactyls from the Dhok Pathan deposits suggest a mosaic of habitats ranging from grassy woodland and woodland, to riverine and forest settings. This is what one would expect in light of the diversity of the animal community and availability of diverse habitat to accommodate it. A similar mosaic palaeoenvironment is inferred for the Hasnot deposits (Khan, 2007). The autecological characterizations of all twenty three species support the reconstruction of an environment mainly shaped by water logged, grassy woodlands, formed under drier conditions and closed vegetation and are thus consistent with current environmental interpretations for the Eurasia and Africa Late Miocene. The artiodactyl community structure of the Dhok Pathan suggest the existence of a vast openland environment with alternating dry and floodseasons which forced a mosaic of ecotonal habitats with an endless number of niches and corresponding adaptations. In summary, the assemblages of the Dhok Pathan type locality represent the late Miocene to Pliocene. The Dhok Pathan type locality fossil assemblages are found in Africa and Eurasia, and they are thus of particular interest to an understanding of mammalian evolutionary processes.
165
REFERENCES
AKHTAR, M., 1992. Taxonomy and Distribution of the Siwalik Bovids. PhD thesis
(unpublished), University of the Punjab, Lahore, Pakistan.
AKHTAR, M., 1995. Pachyportax giganteus, new species (Mammalia, Artiodactyla,
Bovidae) from the Dhok Pathan, district Chakwal, Punjab, Pakistan. Pakistan
Journal of Zoology, 27(4): 337-340.
ALCALÁ, L., MORALES, J., 2006. Antilopinae (Bovidae, Mammalia) from the Lower
Pliocene of Teruel Basin (Spain). Estudios Geológicos, 62(1): 559-570.
ARAMBOURG, C., PIVETEAU, J., 1929. Dorcatherium puyhauberti sp. n. Pontian nr.
Salonica. Ann. Paleont., 18(2-3): 1-34.
ARIF, M., RAZA, S., 1991. New findings of Cervidae (Mammalia) from the Upper
Siwaliks of Potwar-Mirpur Areas, Pakistan. Proceedings Pakistan Congress of
Zoology, 11: 275-281.
ARIF, M., SHAH, S. M., 1991. Cervus rewati sp. nov. (Mammalia, Cervidae) from the
Upper Siwaliks of Pakistan. Pakistan Geological Survey Memoirs, 17: pt.11.
ARIF, M., SHAH, S. M., VOS, J. D., 1991. Cervus triplidens (Mammalia, Cervidae)
from the Upper Siwaliks of Pakistan. Geological Survey of Pakistan Memoirs,
17: pt.11.
AZANZA, B., MORALES, J., 1994. Tethytragus nov. gen et Gentrytragus nov. gen.
Deux nouveaux Bovidés (Artiodactyla, Mammalia) du Miocène moyen.
Proceedings, Koninklijke Nederlandse Akademie Van Wetenschappen, B97:
249-282. 166
AZZAROLI, A.,S 1954. Critical observations upon Siwalik deer. Proceedings of the
Linnaean Society of London, pp. 75-87.
BAKALOV, P., NIKOLOV, I., 1962. Tertiary mammals. In Fossils of Bulgaria 10 (ed.),
V. Tzankov. Sofia: Academie des Sciences de Bulgarie [in Bulgarian].
BARRY, J. C., LINDSAY, E. H., JACOBS, L. L., 1982. A biostratigraphic zonation of
the middle and upper Siwaliks of the Potwar Plateau of northern Pakistan.
Palaeogeography, Palaeoclimatology, Palaeoecology, 37: 95-130.
BARRY, J. C., MORGAN, M. E., WRINKLER, A. J., FLYNN, L. J, LINDSAY, E. H.,
JACOBS, L. L., PILBEAM, D., 1991. Faunal interchange and Miocene
terrestrial vertebrates of southern Asia. Paleobiology, 17: 231-245.
BARRY, J., MORGAN, M., FLYNN, L., PILBEAM, D., BEHRENSMEYER, A. K.,
RAZA, S., KHAN, I., BADGELY, C., HICKS, J., KELLEY, J., 2002. Faunal
and Environmental change in the Late Miocene Siwaliks of Northern Pakistan.
Paleobiology, 28: 1-72.
BARRY, J. C., COTE, S., MACLATCHY, L., LINDSAY, E. H., KITYO, R., RAJPAR,
A. R., 2005. Oligocene and Early Miocene Ruminants (Mammalia,
Artiodactyla) from Pakistan and Uganda, Palaeontologia Electronica, 8(1): 1-
29.
BASU, P. K., 2004. Siwalik mammals of the Jammu Sub-Himalaya, India: an appraisal
of their diversity and habitats. Quaternary International, 117: 105-118.
BEHRENSMEYER, A., BARRY, J., 2005. Biostratigraphic Surveys in the Siwaliks of
Pakistan. A Method for Standardized Surface Sampling of the Vertebrate Fossil
Record. Palaeontologia Electronica, 8 (1): 1-24. 167
BEHRENSMEYER, A. K, WILLIS, B. J, QUADE, J., 1995. Floodplains and paleosols
of Pakistan Neogene and Wyoming Paleogene deposits: a comparative study.
Palaeogeography, Palaeoclimatology, Palaeoecology, 115: 37–60.
BEHRENSMEYER, A. K., 1987. Miocene fluvial facies and vertebrate and vertebrate
taphonomy in northern Pakistan. In recent developments in fluvial
sedimentology (ed. F. G. Ethridge. R. M. Flores, and M. D. Harvey). Society of
Economic Paleontologists and Mineralogists. Special Publication, 39: 169-176.
BEHRENSMEYER, A. K., TAUXE, L., 1982. Isochronous fluvial systems in Miocene
deposits of northern Pakistan. Sedimentology, 29: 331-352.
BERGGERN, W. A., KENT, D. V., FLYNN, L. J. AND VAN COUVERING, J. A.,
1985. Cenozoic geochronology. Geological Society of America Bulletin, 96:
1407-1418.
BERNOR, R. L., 1986. Mammalian biostratigraphy, geochronology, and zoogeographic
relationships of the Late Miocene Maragheh fauna, Iran. Journal of vertebrate
Palaeontology, 6: 76-95.
BHATTI, Z. H., 2005. Taxonomy, Evolutionary History and Biogeography of the
Siwalik Giraffids. PhD thesis (unpublished), University of the Punjab, Lahore,
Pakistan.
BIBI, F., 2007. Origin, Paleoecology, and Paleobiogeography of early Bovini.
Palaeogeography, Palaeoclimatology, Palaeoecology, 248: 60-72.
BIBI, F., GULUC, E., 2008. Bovidae (Mammalia: Artiodactyla) from the Late Miocene
of Sivas, Turkey. Journal of Vertebrate Palaeontology, 28(2): 501–519. 168
BIBI, F., BUKHSIANIDZE, M., GENTRY, A. W., GERAADS, D., KOSTOPOULOS,
D. S., VRBA, E. S., 2009. The Fossil Record and Evolution of Bovidae: State of
the Field. Palaeontologia Electronica, 12(3): 1-11.
BLAINVILLE, H. M., 1816. Sur plusieurs espèces d animaux mammifères de 1’ order
des Ruminants. Bull. Soc. Phil. Paris, pp. 73-82.
BOHLIN, B., 1941. Gazella (Protetraceros) gaudryi (Schlosser) and Gazella
dorcadoides Schlosser. Bulletin Geological Institute, University of Upsala, 28:
79–122.
BONIS, L. DE, KOUFOS, G. D., 1999. The Miocene large mammal succession in
Greece. In: Agusti J., Rook L. & Andrews P. (eds), Hominoid Evolution and
Climatic Change in Europe, volume 1: The Evolution of Neogene Terrestrial
Ecosystems in Europe. Cambridge University Press, Cambridge: 205-237.
BOUVRAIN, G., 1979. Un nouveau genre de Bovide de la fin du Miocene. Bulletin de la
Societe Geologique de France, Paris, 21: 507-511.
BOUVRAIN, G., 1982. Revision du genre Prostrepsiceros Major 1891.
Palaontologische Zeitschrift, 56: 113-124.
BOUVRAIN, G., 1988. Les Tragoportax (Bovidae, Mammalia) des gisements du
Miocène supérieur de Dytiko (Macédonie, Grèce). Annales de Paléontologie,
74(1): 43-63.
BOUVRAIN, G., 1994. Un bovidé du Turolien inférieur d’Europe orientale: Tragoportax
rugosifrons. Annales de Paléontologie, 80(1): 61-87. 169
BOUVRAIN, G., BONIS, L. DE, 1983. Étude d’un Miotragocère du Miocène supérieur
de Macédoine (Grèce). In: Buffetaut É., Mazin J. & Salmon E. (eds), Actes du
symposium paléontologique G. Cuvier. Montbéliard, 1982: 35-51.
BOUVRAIN, G., THOMAS, H., 1992. Une antilope a chevilles spiralees:
Prostrepsiceros zitteli (Bovidae). Miocene superior du Jebel Hamrin en Irak.
Geobios, 25: 525-533.
BOUVRAIN, G., SEN, S., THOMAS, H., 1994. Un nouveau genre d’antilope dans le
Miocène supérieur de Sinap en Turquie. Revue de Paléobiologie 13(2): 375-
380.
BROWN, B., 1926. A new deer from the Siwalik. American Mus. Novitates, 242: 1-6.
CANDE, S. C., KENT, D. V., 1995. Revised calibration of the geomagnetic polarity time
scale for the Cretaceous and Cenozoic. Journal of Geophysical Research, B100:
6093-6095.
CHEN, G. F., 1988. Remarks on the Oioceros species (Bovidae, Artiodactyla,
Mammalia) from the Neogene of China. Vertebrata Palasiatica, 26: 169-172.
CHEN, G., 1997. The genus Gazella Blainville, 1816 (Bovidae, Artiodactyla) from the
Late Neogene of Yushe Basin, Shanxi, Province, China. Vertebrata Palasiatica,
35(4): 233–249.
COLBERT, E. H., 1935. Siwalik mammals in the American Museum of Natural History.
Transactions of the Amererican Philosphical Society, New Series, 26: 1-401.
COOKE, H., WELLS, L., 1951. Fossil Remains from Chelmer, Near Bulawayo, Southern
Rhodesia. South African Journal of Science, 47: 205–209. 170
DENNELL, R. W., COARD, R., TURNER, A., 2008. Predators and Scavengers in Early
Pleistocene southern Asia. Quaternary International, 192: 78-88.
DOWNING, K. F., LINDSAY, E. H., DOWNS, W. R., SPEYER, S. E., 1993.
Lithostratigraphy and vertebrate biostratigraphy of the early Miocene
Himalayan Foreland, Zinda Pir Dome, Pakistan. Sedimentary Geology, 87: 25-
37.
FAROOQ, U., 2006. Studies of Evolutionary Trends in Dentition of the Siwalik
Tragulids. PhD thesis (unpublished), University of the Punjab, Lahore, Pakistan.
FAROOQ, U., KHAN, M. A., AKHTAR, M., KHAN, A. M., 2007a. Dorcatherium
minus from the Siwaliks, Pakistan. Journal of Animal and Plant Sciences, 17(3-
4): 86-89.
FAROOQ, U., KHAN, M. A., AKHTAR, M., KHAN, A. M., 2007b. Dorcabune
anthracotherioides (Artiodactyla, Ruminantia, Tragulidae) from Hasnot, the
Middle Siwaliks, Pakistan. Pakistan Journal of Zoology, 39(6): 353-360.
FAROOQ, U., KHAN, M. A., AKHTAR, M., KHAN, A. M., 2008. Lower Dentition of
Dorcatherium majus (Tragulidae, Mammalia) in the Lower and Middle
Siwaliks (Miocene) of Pakistan. Turkish Journal of Zoology, 32: 91-98.
FLYNN, L. J., 2003. Small mammal indicators of forest paleo-environment in the
Siwalik deposits of the Potwar Plateau, Pakistan. Deinsea, 10: 183-196.
FLYNN, L. J., BARRY, J. C., MORGAN, M. E., PILBEAM, D., JACOBS, L. L.,
LINDSAY, E. H., 1995. Neogene Siwalik Mammalian Lineages: Species
longevities, rates of change, and modes of speciation. Palaeogeography,
Palaeoclimatology, Palaeoecology, 115: 249-264. 171
FLYNN, L. J., QI, G. Q., 1982. Age of the Lufeng, China, hominoid locality. Nature,
298: 746–747.
FRIEDMAN, R., GEE, J., TAUXE, L., DOWNING, K., LINDSAY, E., 1992. The
magnetostratigraphy of the Chitarwata and lower Vihowa formations of the
Dera Ghazi Khan area, Pakistan. Sedimentary Geology, 81: 253-268.
GAUR, R., 1987. Environment and Ecology of Early Man in Northwest India. B.R.
Publishing Corporation, New Delhi.
GENTRY, A., HEINZMANN, E. P. J., 1996. Miocene ruminants of the Central and
Eastern Tethys and Paratethys, in Bernor, R.L., Fahlbusch, V., and Mittmann,
H.W., (eds.), The evolution of Western Eurasian Neogene mammal faunas: New
York, Columbia University Press, 378-391.
GENTRY, A. W., 1966. Fossil Antilopini of East Africa. Bull. Brit. Mus. Nat. Hist.
(Geol.), 12: 45-106.
GENTRY, A. W., 1970. The Bovidae (Mammalia) of the Fort Ternan fossil fauna; pp.
243-323 in L. S. B. Leakey and R. J. G. Savage (eds.), Fossil Vertebrates of
Africa, Vol. 2. Academic Press, London.
GENTRY, A. W., 1971. The earliest goats and other antelopes from the Samos
Hipparion fauna. Bulletin of the British Museum (Natural History), Geology
Series, 20: 229–296.
GENTRY, A. W., 1978. Bovidae; pp. 540-572 in V. J. Maglio and H. B. S. Cooke (eds.),
Evolution of African Mammals. Harvard University Press, Cambridge,
Massachusetts. 172
GENTRY, A. W., 1980. Fossil Bovidae (Mammalia) from Langebaanweg, South Africa.
Annales of South African Museum, 79: 213-337.
GENTRY, A. W., 1985. The Bovidae of the Omo Group deposits, Ethopia. In: Les faunes
Plio-Pleistocenes de la Basse Vallee de l’Omo (Ethopia), tome 1,
Perissodactyles-Artiodactyles (Bovidae), CNRS, Paris, pp. 119-191.
GENTRY, A. W., 1990. Ruminant artiodactyls of Pasalar, Turkey. Journal of Human
Evolution, 19 (4/5): 529-550.
GENTRY, A. W., 1992. The subfamilies and tribes of Bovidae. Mammal Review, 22:
1-32.
GENTRY, A. W., 1994. The Miocene differentiation of Old World Pecora (Mammalia).
Historical Biology, 7: 115-158.
GENTRY, A. W., 1997. Fossil ruminants (Mammalia) from the Manonga Valley,
Tanzania. In Neogene Paleontlogy of the Manonga Valley, Tanzania, pp. 107-
135 (ed. T. J. Harrison). Plenum Press, New York.
GENTRY, A. W., 1999. Fossil Pecorans from the Baynunah Formation, Emirate of Abu
Dhabi, United Arab Emirates. Chap. 22 in Abu Dhabi pecorans, Yale University
Press, New Haven, 290-316.
GENTRY, A. W., 2003. Ruminantia (Artiodactyla). In: Fortelius, M., Kappelman, J.,
Sen, S., Bernor, R. (eds.). Geology and Paleontology of the Miocene Sinap
Formation, Turkey. – 332– 379, Columbia University Press, New York.
GENTRY, A. W., ROSSNER, G. E., HEIZMANN, E. P. S., 1999. Suborder Ruminantia.
In Rossner, G.E., and Heissig, K. (eds.), The Miocene land mammals of Europe:
Munchen, Verlag Dr. Friedrich Pfeil, 225-258. 173
GERAADS, D., SPASSOV, N., KOVACHEV, D., 2003. Palaeoreas lindermayeri
(Wagner, 1848) from the upper Miocene of Bulgaria, and a revision of the
species. Geodiversitas, 25(2): 405-415.
GERAADS, D., 1989. Vertebres fossils du Miocene superieur du Djebel Krechem el
Artsouma (Tunisie centrale): Comparisons biostratigraphiques. Geobios, 22:
777-801.
GHAFFAR, A., 2005. Studies on equids, cervids and Carnivora from the Siwalik Hills of
Pakistan. PhD thesis (unpublished), University of the Punjab, Lahore, Pakistan.
GHAFFAR, A., AKHTAR, M., KHAN, M. A., NAZIR, M., 2004. Report on Cervus
sivalensis from the Upper Siwaliks of Pakistan. Punjab Univ. J. Zool., 19: 83-
88.
GROMOLARD, C., 1980. Une nouvelle interpretation des grands Bovidae (Artiodactyla,
Mammalia) du Pliocene d’Europe occidentale classes jusqu’a present dans le
genre Parabos: Parabos cordieri (de Christol) emend., ?Parabos boodon
(Gervais) et Alephis lyrix n. gen., n. sp. Geobios, 13: 767-775.
GUO, J., DAWSON, M. R., BEARD, K. C., 2000. Zhailimeryx, a new lophiomerycid
artiodactyl (Mammalia) from the late middle Eocene of central China and the
early evolution of ruminants. Journal of Mammalian Evolution, 7: 239-258.
HAILE-SELASSIE, Y., WOLDE GABRIEL, G., WHITE, T., BERNOR, R. L.,
DEGUSTA, D., RENNE, P. R., HART, W. K., VRBA, E. S., STANLEY, A.,
AND HOWELL, F. C. 2004. Mio-Pliocene mammals from the Middle Awash,
Ethiopia. Géobios, 37: 536-552. 174
HAN, DE-FEN., 1974. First discovery of Dorcabune in China. Vertebrate Palasiatica,
12(3): 217-221.
HEINTZ, E., 1971. Gazella deperdita (Gervais) 1847 (Bovidae, Artiodactyla,
Mammalia) du Pontien du Mont Lubéron, Vaucluse, France. Annales de
Paléontologie, 57: 209–229.
HEISSIG, K., 1972. Palaontologische und geologische untersuchungen im tertiar von
Pakistan. 5. Rhinocerotidae(Mamm.) aus den unteren und mittleren Siwalik-
Schichten. Abh. Bayer. Akad. Wiss., Munich (NF), 152: 1-112.
IQBAL, M., LIAQAT, A., KHAN, M. A., AKHTAR, M., 2009. Some New Remains of
Hipparion from the Dhok Pathan Type Locality, Pakistan. Journal of Animal and
Plant Sciences, 19(3): 154-157.
JANIS, C. M., DAMUTH, J., THEODOR, J. M., 2002. The origins and evolution of the
North American grassland biome: the story from the hoofed mammals.
Palaeogeography, Palaeoclimatology, Palaeoecology, 177: 183–198.
JOHNSON, G. D., ZEITLER, P., NAESER, C. W., JOHNSON, N. M., SUMMERS, D.
M., FROST, C. D., OPDYKE, N. D. AND TAHIRKHELI, R. A., 1982. The
occurrence and fission-track ages of Late Neogene and Quaternary volcanic
sediments, Siwalik group, northern Pakistan. Palaeogeography,
Palaeoclimatology, Palaeoecology, 37: 63-93.
JOHNSON, N. M., OPDYKE, N. D., JOHNSON, G. D., LINDSAY, E. H. AND
TAHIRKHELI, R. A., 1982. Magnetic polarity, stratigraphy and ages of Siwalik
group rocks of the Potwar Plateau, Pakistan. Palaeogeography,
Palaeoclimatology, Palaeoecology, 37: 17-42. 175
JOHNSON, N. M., STIX, J., TAUXE, L., CERVENY, P. F. AND TAHIRKHELI, R. A.,
1985. Paleomagnetic chronology, fluvial processes and tectonic implications of
the Siwalik deposits near Chinji Village, Pakistan. Journal of Geology, 93: 27-
40.
KAHLKE, H. D., 1976. Die Cervidenreste aus den Travertinen von Taubach.
Quartarpalaontologie, 2: 209–223.
KAPPELMAN, J. W., 1986. The paleoecology and chronology of the Middle Miocene
hominoids from the Chinji formation of Pakistan. PhD thesis, Harvard
University.
KHAN, A. M., 2010. Taxonomy and Distribution of Rhinoceros from the Siwalik Hills of
Pakistan PhD thesis (unpublished), University of the Punjab, Lahore, Pakistan.
KHAN, M. A., 2007. Taxonomic Studies on Fossil Remains of Ruminants from Tertiary
Hills of Hasnot, Pakistan. PhD thesis (unpublished), University of the Punjab,
Lahore, Pakistan.
KHAN, M. A., GHAFFAR, A., FAROOQ, U., AKHTAR, M., 2007. New Fossil remains
of Selenoportax vexillarius from the Late Miocene of Hasnot. Pakistan
Journal of Zoology, 39(5): 333-338.
KHAN, M. A., 2008. Fossil bovids from the late Miocene of Padri, Jhelum, Pakistan.
Pakistan Journal of Zoology, 40(1): 25-29.
KHAN, M. A., IQBAL, M., GHAFFAR, A., KHAN, A. M., AKHTAR, M. 2008a.
Collection of Selenoportax from the Kundi Wala Kas, Hasnot, Pakistan.
Pakistan Journal of Zoology, 40(4): 303-307. 176
KHAN, M. A., AHMAD, Z., AKHTAR, M., 2008b. Selenoportax vexillarius from Dhok
Pathan Formation of the Middle Siwaliks, Pakistan. Biologia (Pakistan), 54(1):
65-72.
KHAN, M. A., ILIOPOULOS, G., AKHTAR, M., 2009a. Boselaphines (Artiodactyla,
Ruminantia, Bovidae) from the Middle Siwaliks of the Hasnot, Pakistan.
Geobios, 42: 739-753.
KHAN, M. A., IQBAL, M., AKHTAR, M., HASSAN, M., 2009b. Chalicotheres in the
Siwaliks of Pakistan. Pakistan Journal of Zoology, 41(6): 429-435.
KOHLER, M., 1987. Boviden des turkischen Miozans (Kanozoikum und Braunkohlen
der Turkei 28). Paleontologia Evolucio, 21: 133-246.
KOHLER, M., 1993. Skeleton and habitat of recent and fossil ruminants. Munchner
Geowiss. Abh. (A) 25, 88p.
KOSTOPOULOS, D. S., 2005. The Bovidae (Mammalia, Artiodactyla) from the late
Miocene of Akkasdagı, Turkey. Geodiversitas, 27 (4): 747–791.
KOSTOPOULOS, D. S., 2006. Late Miocene Vertebrate Locality of Perivolaky,
Thessaly, Greece. Palaeontographica Abt., 276: 151–183.
KOSTOPOULOS, D. S., 2009. The late Miocene Mammal Faunas of the Mytilinii Basin,
Samos Island, Greece: New Collection. 14. Bovidae. Beitr. Paläont. 31: 339–
383.
KOSTOPOULOS, D. S., KOUFOS, G. D., 1996. Late Miocene bovids (Mammalia,
Artiodactyla) from the locality “Nikiti-1” (NKT), Macedonia, Greece. Annales
de Palantologie, 81: 251–300. 177
KRETZOI, M. 1968. New generic names for homonyms. Vertebrata Hungarica, 10(1-2):
163-166.
LEHMANN, U., THOMAS, H., 1987. Fossil Bovidae (Mammalia) from the Mio-
Pliocene of Sahabi (Libya), in BOAZ N. T., ARNAUTI E. L., GAZIRY A. W.
& HEINZELIN J. DE (eds), Neogene Paleontology and Geology of Sahabi,
Libya. Liss, New York, pp. 323-335.
LISTER, A. M., 1984. Evolutionary and Ecological Origins of British Deer. Proceedings
of the Royal Society of Edinburgh, 82B: 205–229.
LISTER, A. M., 1986. New Results on Deer from Swanscombe, and the Stratigraphical
Significance of Deer Remains in the Middle and Upper Pleistocene of Europe.
Journal of Archaeological Science, 13: 319–338.
LYDEKKER, R., 1876. Molar teeth and other remains of Mammalia from the India
Tertiaries. Pal. Ind., 10(2): 19-87.
LYDEKKER, R., 1878. Indian Tertiary and Post-Tertiary vertebrate. 3. Crania of
Ruminants. Palaeontologia Indica, 10: 88-181.
LYDEKKER, R., 1880. A sketch of the history of the fossil vertebrata of India. Journal
Asiatic Society of Bengal, XLIX: 8-40, Pt.2.
LYDEKKER, R., 1884. Additional Siwalik Perissodactyla and Proboscidea. Memoirs of
the Geological Survey of India. Palaeontologica Indica, 3(10): 1–34.
MATTHEW, W. D., 1929. Critical observations upon Siwalik Mammals. Bulletin of
American Museum of Natural History, 56: 435-560.
MERCERON, G., ZAZZO, A., SPASSOV, N., GERAADS, D., KOVACHEV, D., 2006.
Bovid Palaeoecology and Palaeoenvironments from the Late Miocene of 178
Bulgaria: Evidence from dental microwear and stable isotopes. Palaeogeography,
Palaeoclimatology, Palaeoecology, 241(3-4): 637–654.
MÉTAIS, G., CHAIMANEE, Y., JAEGER, J., DUCROCQ, S., 2001. New remains of
primitive ruminants from Thailand: evidence of the early evolution of the
Ruminantia in Asia. Zoologica Scripta, 30: 231–248.
METAIS, G., GUO, J., BEARD, K. C., 2004. A New Small Dichobunid Artiodactyl from
Shanghuang (Middle Eocene, Eastern China): Implications for the Early
Evolution of Proto-Selenodonts in Asia. Bulletin of Carnegie Museum of
Natural History, 36: 177-197.
MOYA-SOLA , S., 1988. Morphology of Lower Molars of the Ruminants (Artiodactyla,
Mammalia): Phylogenetic Implication. Paleontol. Evoluc., 22: 61–70.
MOYA-SOLA, S., 1983. Les Boselaphini (Bovidae, Mammalia) del Neogeno de la
Peninsula Iberica. Dissertation, Universidad Autonoma de Barcelona,
Publicaciones de Geologia, 18: 1-236.
MOYA-SOLA, S., 1983. Los Boselaphini (Bovidae, Mammalia) del Neogeno de la
Peninsula Iberica. Dissertation, Universidad Autonoma de Barcelona,
Publicaciones de Geologia, 18: 1-236.
NAKAYA, H., 1994. Faunal change of Late Miocene Africa and Eurasia: mammalian
fauna from the Namurungule Formation, Samburu Hills, northern Kenya.
African study Monographs. Supplementary Issue, 20: 1–112.
NANDA, A. C., 2008. Comments on the Pinjor Mammalian Fauna of the Siwalik Group
in relation to the Post-Siwalik Faunas of Peninsular India and Indo-Gangetic
Plain. Quaternary International, 192: 6-13. 179
OPDYKE, N. D., LINDSAY, E., JOHNSON, G. D., JOHNSON, N., TAHIRKHELI, R.
A., MIRZA, M. A., 1979. Magnetic polarity, stratigraphy and vertebrate
paleontology of the Upper Siwalik Subgroup of northern Pakistan.
Palaeogeography, Palaeoclimatology, Palaeoecology, 27: 1-34.
PICKFORD, M., 1988. Revision of the Miocene Suidae of the Indian Subcontinent.
Münchner Geowissenschaften Abhandlungen, 12: 1-91.
PICKFORD, M., 2001. Africa’s smallest ruminant: A new tragulid from the Miocene of
Kenya and the biostratigraphy of East African Tragulidae. Geobios, 34(4): 437–
447.
PICKFORD, M., 2002. Ruminants from the Early Miocene of Napak, Uganda. Annales
de Paléontologie, 88: 85−113.
PICKFORD, M., SENUT, B., MOURER-CHAUVIRE, C., 2004. Early Pliocene
Tragulidae and peafowls in the Rift Valley, Kenya: Evidence for rainforest in East
Africa. Comptes Rendus Palevol, 3: 179–189.
PILBEAM, D., BARRY, J., MEYER, G. E., SHAH, S. M. I., PICKFORD, M. H. L.,
BISHOP, W. W., THOMAS, H., JACOBS, L. L., 1977. Geology and
palaeontology of Neogene strata of Pakistan. Nature, 270: 684-689.
PILGRIM, G. E., 1910. Notices of new Mammalian genera and species from the
Tertieries of India-Calcutta. Records of the Geological Survey of India, 40: 63-
71.
PILGRIM, G. E., 1911. The fossil Giraffidae of India. Memior of the Geological Survey
of India. Palaeontologica Indica, New Series, 4 (1): 1-29. 180
PILGRIM, G. E., 1913. Correlation of the Siwaliks with Mammal Horizons of Europe.
Records of the Geological Survey of India, 43(4): 264-326.
PILGRIM, G. E., 1915. The dentition of the Tragulid genus Dorcabune. Records of the
Geological Survey of India, 45: 226–238.
PILGRIM, G. E., 1926a. The Tertiary formations of India and the interrelation of marine
and terrestrial deposits. Proceedings of Pan-Pacific Science Congress,
Australia, pp. 896-931.
PILGRIM, G. E., 1926b. The fossil Suidae of India. Memoirs of the Geological Survey of
India, New Series, 8(4): 1-65.
PILGRIM, G. E., 1937. Siwalik antelopes and oxen in the American Museum of Natural
History. Bulletin of American Museum of Natural History, 72: 729-874.
PILGRIM, G. E., 1939. The fossil Bovidae of India. Pal. Ind., N .S., 26(1): 1-356.
PILGRIM, G. E., HOPWOOD, A. T., 1928. Catalogue of the Pontian Bovidae of Europe
in the Department of Geology. British Museum (Natural History), London.
PITRA, C., FICKELA, J., MEIJAARD, E., GROVES, C. P. 2004. Evolution and
phylogeny of old world deer. Molecular Phylogenetics and Evolution, 33: 880–
895.
QIU, Z. X., QIU, Z. D., 1995. Chronological sequence and subdivision of Chinese
Neogene mammalian faunas. Palaeogeography, Palaeoclimatology,
Palaeoecology, 116: 41–70.
QIU, Z., GU, Y., 1991. The Aragonian vertebrate fauna of Xiacaowan-8. Dorcatherium
(Tragulidae, Artiodactyla). Vertebrata Palasiatica, 29(1): 21–37. [Chinese 21–31;
English 32–37]. 181
QUADE, J., CERLING, T. E., 1995. Expansion of C4 grasses in the Late Miocene of
Northern Pakistan: evidence from stable isotopes in paleosoles.
Palaeogeography, Palaeoclimatology, Palaeoecology, 115: 91 –116.
RAZA, S. M., CHEEMA, I. U., DOWNS, W. R., RAJPAR, A. R. & WARD, S. C., 2002.
Miocene stratigraphy and mammal fauna from the Sulaiman Range,
Southwestern Himalayas, Pakistan. Palaeogeography, Palaeoclimatology,
Palaeoecology, 186:185 –197
ROMER, A. S., 1974. Vertebrate palaeontology, III. The University of Chicago Press,
Chicago Illinois, pp.1-687.
ROSSNER, G. E., MORS, T., 2001. A new find of the enigmatic Eurasian Miocene
ruminant artiodactyls Orygotherium: its meaning for stratigraphic setting and
palaeoecological interpretations. Journal of Vertebrate Paleontology, 21(3):
591-595.
ROSSNER, G. E., 2010. Systematics and palaeoecology of Ruminantia (Artiodactyla,
Mammalia) from the Miocene of Sandelzhausen (southern Germany, Northern
Alpine Foreland Basin). Palaontolology Zurich, 1-40. (DOI 10.1007/s12542-
010-0052-2).
SACH, V., HEIZMANN, E. P. J., 2001. Stratigraphie und Sa¨ugetierfaunen der
Brackwassermolasse in der Umgebung von Ulm (Sudwestdeutschland).
Stuttgarter Beitrage zur Naturkunde, B310: 1–95.
SAVAGE, D. E., RUSSEL, D. E., 1983. Mammalian Paleofaunas of the World, London.
Addison-Wesley, pp. 1-432. 182
SENYUREK, M. S., 1954. A Horn-core of Gazella capricornis Rodler and Weithofer
found at Kucukyozgat. Belleten, XVIII: 142–145.
SHAH, S. M. I., 1977. Stratigraphy of Pakistan. Memoirs of the Geological Survey of
Pakistan, 12: 1-138.
SHAH, S. M. I., 1980. Stratigraphy and economic geology of Central Salt Range.
Records of the Geological Survey of Pakistan, 52: 1-104.
SOLOUNIAS, N., 1981. The Turolian fauna from the Island of Samos, Greece.
Contribution to Vertebrate Evolution, 6: 99-232.
SOLOUNIAS, N., BARRY, J. C., BERNOR, R. L., LINDSAY, E. H., RAZA, S. M.,
1995. The Oldest Bovid from the Siwaliks, Pakistan. Journal of Vertebrate
Paleontology, 15 (4): 806-814.
SOMMER, R. S., ZACHOS, F. E., STREET, M., JORIS, O., SKOG, A., BENECKE,
N., 2008. Late Quaternary Distribution Dynamics and Phylogeography of the
Red Deer (Cervus elaphus) in Europe. Quaternary Science Reviews, 27: 714-
733.
SPASSOV, N., GERAADS, D., 2004. Tragoportax Pilgrim, 1937 and Miotragocerus
Stromer, 1928 (Mammalia, Bovidae) from the Turolian of Hadjidimovo,
Bulgaria, and a revision of the late Miocene Mediterranean Boselaphini.
Geodiversitas, 26(2): 339–370.
STEININGER, F. F., 1999. Chronostratigraphy, geochronology and biochronology of the
Miocene “European land mammal mega-zones” (ELMMZ) and the Miocene
“mammal-zones” (MN-zones). In: Rössner Gertrud, E., Heissig, K., Fahlbusch, 183
V. (Eds.), The Miocene Land Mammals of Europe. Verlag Friedrich Pfeil,
Munich, pp. 9–24.
STIX, J., 1982. Stratigraphy of the Kamlial Formation near Chinji Village, northern
Pakistan. BA thesis, Dartmouth College, Hanover, N. H.
SURESH, N., SUMIT, K., ROHTASH, G. K., SANGODE, S. J., 2004. Clay-
mineral distribution patterns in late Neogene fluvial sediments of the Subathu
sub-basin, central sector of Himalayan foreland basin: implications for
provenance and climate. Sedimentary Geology, 163: 265–278.
TAUXE, L. AND OPDYKE, N. D., 1982. A time framework based on
magnetostratigraphy for the Siwalik sediments of the Khaur area, northern
Pakistan. Palaeogeography, Palaeoclimatology, Palaeoecology, 37: 43-61.
TEDFORD, R. H., FLYNN, L. J., ZHANXIANG, Q., OPDYKE, N. D., DOWNS, W. R.,
1991. Yushe Basin, China: Paleomagnetically calibrated mammalian
biostratigraphic standard for the late Neogene of eastern Asia. Journal of
Vertebrate Paleontology, 11: 519–526.
THOMAS, H., 1979. Les Bovides miocenes des rifts est-africains: Implications
paleobiogeogarphiques. Bulletin Geological Society of France, 7e Series,
XXI(3): 295-299.
THOMAS, H., 1980. Les bovides du Miocene superieur des couches de Mpesida et de la
formation de Lukeino (district de Baringo, Kenya), in: Proceedings of the 8th
Pan-African Congress of Prehistory, Nairobi, 1977 (R. E. F. Leakey and B. A.
Ogot, eds.), Nairobi, pp. 82-91. 184
THOMAS, H., 1981. Les bovides miocenes de la formation de Ngorora du Basin de
Baringo (Kenya), Proc. K. Nederl. Akad. Wetensch. B84: 335-409.
THOMAS, H., 1984. Les bovides ante-hipparions des Siwaliks inferieurs (plateau du
Potwar), Pakistan. Memoires de la Societe Geologique de France, Paris, 145:
1-68.
VRBA, E. S., 1995. The fossil record of African antelopes (Mammalia, Bovidae) in
relation to human evolution and palaeoclimate. In Paleoclimate and Evolution:
with emphasis on human origins (eds. E. S. Vrba, G. H. Denton, T. C. Partridge
and L. H. Burkle), pp. 385–424. Yale University Press, New Haven.
VRBA, E. S., HAILE-SELASSIE, Y., 2006. A New Antelope, Zephyreduncinus
oundagaisus (Reduncini, Artiodactyla, Bovidae), from the Late Miocene of the
Middle Awash, Afar Rift, Ethiopia. Journal of Vertebrate Paleontology, 26(1):
213 –218.
VRBA, E. S., SCHALLER, G., 2000. Phylogeny of Bovidae based on behavior, glands,
skulls, and postcrania, p. 203-222. In Vrba, E.S., and Schaller, G. (eds.),
Antelopes, Deer, and Relatives. Yale University Press, New Haven.
WEST, R. M., 1980. A minute new species of Dorcatherium (Tragulidae, Mammalia)
from the Chinji Formation near Daud Khel, Mianwali district, Pakistan. MILW.
Public Mus. Contribution to Biology and Geology, 3(33): 1-6.
WHITWORTH, T., 1958. Miocene ruminants of East Africa. Fossil Mammals of Africa.
Bulletin of British Museum ofNatural History, 15: 1-50. 185
WILLIS, B. J., BEHRENSMEYER A. K., 1995. Fluvial systems in the Siwalik Neogene
and Wyoming Paleogene. Palaeogeography, Palaeoclimatology,
Palaeoecology, 114: 13-35.
WILLIS, B., 1993a. Ancient river systems in the Himalayan foredeep, Chinji village
area, northern Pakistan. Sedimentary Geology, 88: 1-76.
WILLIS, B., 1993b. Evolution of Miocene fluvial systems in the Himalayan foredeep
through a two kilometer thick succession in northern Pakistan. Sedimentary
Geology, 88: 77-121.
186
APPENDICES
Appendix 1 List of the studied artiodactyls from Dhok Pathan.
Artiodactyla Bovidae Tragoportax T. punjabicus T. browni T. cf. salmontanus Miotragocerus M. large sp. M. cf. gluten Pachyportax P. latidens cf. P. nagrii Selenoportax S. cf. vexillarius Prostrepsiceros P. cf. vinayaki Gazella G. lydekkeri Kobus K. porrecticornis cf. Kobikeryx sp. Caprotragoides C. cf. potwaricus Cervidae Cervus C. cf. sivalensis C. cf. triplidens Tragulidae Dorcatherium D. majus Dorcabune D. anthracotherioides D. cf. nagrii Giraffidae Hydaspitherium H. cf. megacephalum H. cf. grande H. sp. Suidae Propotamochoerus P. hysudricus Hippopotamodon ?H. sp. 187
Appendix 2
Studied material
Tragoportax punjabicus
Skull: PUPC 86/149, opisthocranium. PUPC 86/150, opisthocranium. Horn-core: PUPC
86/264, broken at the apex, left side. Upper dentition: PUPC 83/743, isolated right P3.
PUPC 87/327, isolated left P3. PUPC 00/21, isolated left P3. PUPC 00/89, right P4-M1.
PUPC 83/671, isolated right M1. PUPC 87/677, isolated right M1. PUPC 83/673, isolated
left M1. PUPC 83/30, isolated left M1. PUPC 83/676, isolated right M1. PUPC 83/674,
isolated right M1. PUPC 97/13, isolated right M1. PUPC 00/80, right M1-2. PUPC
85/242, isolated right M2. PUPC 97/06, isolated left M2. PUPC 83/603, isolated right
M2. PUPC 83/10, isolated left M2. PUPC 83/666, isolated right M2. PUPC 00/01, isolated left M2. PUPC 83/693, isolated right M2. PUPC 08/126, palatine parts of both maxillae with right P3-M3 and left P2-M3. PUPC 83/275, right maxillary fragment with
P3-M3. PUPC 97/10, right maxillary fragment with M1-M3. PUPC 83/665, isolated left
M3. PUPC 97/7, isolated right M3. PUPC 83/688, isolated right M3. Lower dentition:
PUPC 83/712, isolated left p2. PUPC 97/12, left mandible fragment with p2-p4. PUPC
83/204, isolated left p3. PUPC 87/324, isolated left p3. PUPC 83/711, isolated left p4.
PUPC 68/237, isolated left p4. PUPC 97/3, isolated right m1. PUPC 87/282, isolated left m1. PUPC 86/26, isolated left m1. PUPC 83/700, isolated right m1. PUPC 86/25, isolated left m2. PUPC 83/617, isolated left m2. PUPC 99/28, isolated right m2. PUPC 83/701, isolated left m2. PUPC 83/680, isolated left m3. PUPC 83/767, isolated left m3. PUPC
83/277, left mandible fragment with m3.
188
Tragoportax browni
Opisthocranium: PUPC 86/204. Horn-cores: PC-GCUF 09/11, almost complete, side
indeterminate. PUPC 87/1, fragment, side indeterminate. Upper dentition: PUPC 87/281,
isolated left P3. PC-GCUF 09/42, isolated right P3. PC-GCUF 09/43, isolated right P4.
PUPC 83/689, isolated left M1. PUPC 86/151, left maxillary fragment with M1-M2.
PUPC 86/29, left maxillary fragment with M1-M2. PUPC 83/861, left maxillary fragment with M1-M2. PUPC 97/80, isolated left M2. PUPC 83/662, isolated right M2. PUPC
87/324, right maxillary fragment with M2-M3. PUPC 83/283, right maxillary fragment
with M2-M3. Lower dentition: PUPC 99/27, right mandible fragment with m1-m2. PUPC
83/265, left mandible fragment with m2-m3. PUPC 83/269, right mandible fragment with
m2-m3. PUPC 97/11, left mandible fragment with m3.
Tragoportax cf. salmontanus
Horn-core: PUPC 93/20, incomplete, left side.
Miotragocerus large sp.
Upper dentition: PUPC 83/275, isolated right P2. PUPC 86/33, isolated right P3. PUPC
86/39, isolated left P3. PUPC 09/44, right maxillary ramus with P2-P4. PUPC 83/209,
left maxillary ramus with P2-M1. PUPC 87/02 and PUPC 86/308, right maxillary ramus
with DP4-M1. PUPC 87/246, left maxillary ramus with DP3-M2. PUPC 87/115, left
maxillary ramus with DP4-M2. PUPC 87/116 and PUPC 87/117, right and left maxillary
rami with M1-M2. Lower dentition: PUPC 88/740, left mandibular ramus with p2. PUPC
83/703, isolated left p2. PUPC 83/708 and PUPC 83/709, isolated right and left p3s.
PUPC 83/742 and PUPC 87/294, isolated right and left p4s. PUPC 93/277, left
mandibular ramus with p4-m1. PUPC 09/86, left mandibular ramus with p4-m3. PUPC 189
96/05, left mandibular ramus with p4-m1. PUPC 83/277, right mandibular ramus with m2 and broken m3.
Miotragocerus cf. gluten
Horn-core: PUPC 01/01, incomplete, left side.
Pachyportax latidens
Horn-core: PUPC 96/18, basal fragment, right side. Upper dentition: PUPC 80/646, isolated M2. PUPC 80/203, isolated right M2. PUPC 09/74, isolated M2. Lower dentition: PUPC 08/87, right mandibular ramus with m3 with broken hypoconulid and an anterior part of m2. cf. Pachyportax nagrii
Upper dentition: PUPC 99/50, partially damaged isolated ?M3.
Selenoportax cf. vexillarius
Lower dentition: PUPC 99/47, right mandibular ramus with m2 and roots of m1. PUPC
99/22, isolated left semierupted molar. PUPC 00/45, isolated left m2. PUPC 07/16, partially damaged molar. PUPC 97/02, isolated right m3.
Prostrepsiceros cf. vinayaki
Horn-cores: PUPC 87/319, almost complete, left side. PUPC 87/318, incomplete, probably right side.
Gazella lydekkeri
Horn-cores: PUPC 83/319, almost complete with a piece of frontal, broken at the apex, left side. PC-GCUF 09/45, almost complete with supraorbital pit, left side. Lower dentition: PUPC 08/136, right isolated m2. PUPC 07/13 and PUPC 09/76, isolated left m3s. 190
Kobus porrecticornis
Horn-cores: PUPC 87/124 A+B, left and right horn-cores, proximally preserved
fragments, perhaps same individual. PUPC 02/126 and PUPC 87/180, partial left horn-
cores.
cf. Kobikeryx sp.
Horn-cores: PUPC 99/49, distally preserved fragments, side indeterminate.
Caprotragoides cf. potwaricus
Horn-core: PC-GCUF 09/44, right side.
Cervus cf. sivalensis
Lower dentition: PUPC 04/18, right mandibular ramus with m2-3. PUPC 09/73, right
mandibular ramus with m2.
Cervus cf. triplidens
Upper dentition: PUPC 09/71, right maxillary ramus with P4-M2.
Dorcatherium majus
Upper dentition: PUPC 03/14, isolated left M2. Lower dentition: PUPC 01/21, isolated
right m1. PUPC 01/26, left mandibular ramus with m2 and posterior portion of m1.
PUPC 07/09, isolated left m2.
Dorcabune anthracotherioides
Upper dentition: PUPC 99/338, isolated right M2. Lower dentition: PUPC 04/01, right
mandibular ramus with m3.
Dorcabune cf. nagrii
Upper dentition: PUPC 96/45, an isolated left probably M1.
191
Hydaspitherium cf. megacephalum
Lower dentition: PUPC 09/78, left mandibular ramus with broken m1, complete m2 and
partially erupted m3.
Hydaspitherium cf. grande
Lower dentition: PUPC 09/77, right mandibular ramus with m2-3.
?Hydaspitherium sp.
Lower dentition: PUPC 83/267, right mandibular ramus with m2-3.
Propotamochoerus hysudricus
Lower dentition: PUPC 07/32, left mandibular ramus with p4-m3; the p3 roots are also preserved in the alveoli. PUPC 99/02, left mandibular ramus with p4-m1. PUPC 94/65, right mandibular ramus with p4-m2. PUPC 97/15, right isolated m3.
?Hippopotamodon sp.
Lower dentition: PUPC 00/14, left mandibular ramus with broken p4 and partially
complete m1. PUPC 08/26, badly crushed right mandibular ramus with damaged m1-3,
probably juvenile animal.