Bramatherium (Artiodactyla, Ruminantia, Giraffidae) from the Middle Siwaliks of Hasnot, Pakistan: Biostratigraphy and Palaeoecology
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Turkish Journal of Earth Sciences Turkish J Earth Sci (2014) 23: 308-320 http://journals.tubitak.gov.tr/earth/ © TÜBİTAK Research Article doi:10.3906/yer-1112-11 Bramatherium (Artiodactyla, Ruminantia, Giraffidae) from the Middle Siwaliks of Hasnot, Pakistan: biostratigraphy and palaeoecology Muhammad Akbar KHAN*, Muhammad AKHTAR, Ammara IRUM Palaeontology Laboratory, Department of Zoology, Quaid-e-Azam Campus, University of the Punjab, Lahore, Punjab, Pakistan Received: 28.12.2011 Accepted: 10.10.2013 Published Online: 21.03.2014 Printed: 18.04.2014 Abstract: We describe here new remains of the giraffid Bramatherium, from the late Miocene-early Pliocene of Hasnot, northern Pakistan. The fossil giraffid material from Hasnot represents 2 taxa of Bramatherium, B. megacephalum and B. grande. The Hasnot locality is characterised by high proportions of ruminants, namely Boselaphini, Antilopini, Sivatheriinae, Cervini, Dorcatherium, and Dorcabune. The Hasnot fossil ruminant assemblage is fairly typical for the Siwalik province and comparisons with the Greco- Iranian-Afghan and African palaeobiological provinces indicate a late Miocene-early Pliocene age. It is suggested that local Hasnot palaeoenvironments comprised wetlands to woodland biomes, perhaps devoid of expansive dense forests. Key words: Vertebrates, Mammalia, Artiodactyla, late Miocene, Siwaliks, Hasnot 1. Introduction certainly the age of gigantic giraffids. Many species of the Giraffoids with ossicones were first found in the early sivatheriine giraffes are known from the beds of this age Middle Miocene in Africa or their likely ancestors were (Khan et al. 2010). The Tatrot Formation (ca. 3.4–2.6 Ma) recorded in the Middle Miocene at Bugti in Pakistan (Barry is still without any record of the giraffids (Bhatti 2004). The et al. 2005). Giraffid history started in the Early Miocene Pinjor Formation (ca. 2.5–0.6 Ma) is characterised by the of Zebel Zelten in northern Africa with Zarafa Hamilton, presence of Sivatherium giganteum Falconer and Cautley, 1973 and Prolibytherium Arambourg, 1961. Giraffids have 1836 and Giraffa punjabiensis Pilgrim, 1911 (Nanda 2002; a fairly rich fossil record; approximately 30 species have Dennell et al. 2006). The primary goal of this paper is to been found throughout the Neogene of the Old World report new fossil material of Bramatherium from Hasnot (Bohlin 1926; Hamilton 1978; Heintz et al. 1981; Geraads in northern Pakistan (Figure 1) and discuss a provisional 1985, 1986; Janis & Scott 1987; Gentry & Hooker 1988; synonymy of Hydaspitherium Lydekker, 1876 with Gentry 1993; Geraads & Güleç 1999; Geraads et al. 2004, Bramatherium Falconer, 1845, as suggested by Geraads 2005; Kostopoulos & Saraç 2005; Khan & Akhtar 2005– & Güleç (1999) in the Greco-Iranian province. Based on 2006). Today, the family Giraffidae is represented by only the recorded ruminant faunal elements from Hasnot, the 2 species, the okapi [Okapia johnstoni (Sclater, 1901)] and biostratigraphy and palaeoecology of this area are also the giraffe [Giraffa camelopardalis (Linnaeus, 1758)]. Both discussed. modern species are found in Africa. The okapi is present 1.1. Locality in a limited area of heavy forests in West Africa, whereas Hasnot is situated in the west of Jhelum district (32°49ʹN, the giraffe is found in the more open parts of Sub-Saharan 73°18ʹE), Punjab, northern Pakistan (Figure 1A). The Africa (Solounias 2007; Harris et al. 2010). locality of Hasnot yielded a rich mammalian fauna (Table In the Siwaliks (Figure 1A – chronostratigraphic 1), which provides the opportunity for high-resolution context), the giraffids made their first appearance in the sampling and reconstruction of palaeoenvironments lower portion of the Chinji Formation (Matthew 1929; (Pilbeam et al. 1977; Barry et al. 2002; Bhatti 2004; Farooq Colbert 1935; Bhatti 2004). The large or sivatheriine- et al. 2007a, 2007b, 2007c, 2007d, 2008; Khan 2007, 2008; sized giraffids were not present in the Chinji Formation. Khan et al. 2007, 2008, 2009, 2010; Ghaffar et al. 2010, As indicated by the fossil record, the sivatheriines were 2011). The fossiliferous localities around the village definitely present in the Nagri Formation (Colbert of Hasnot are stratigraphically situated in 3 different 1935; Bhatti 2004). The Dhok Pathan Formation was formations (Figure 1B), and they have yielded fossil * Correspondence: [email protected] 308 KHAN et al. / Turkish J Earth Sci 72 o 73oo74 Ma e Formations Ma A 0 Boulder Islalammabaabad Conglomerates 0.6 PoPottwwarar Plateaeauu s du Pleistocen r In Pinjor an N 2 pe PAKISISTTAAN So Up 2.6 Tatrot e 3.4 Karachrachii 4 cen o 34 Plio Margala Hills r Ka an la Ch th itta H 6 s Rive ill Islamabad s Pa du In Middle 8 Dhok p Kheur ou u 10 lik Gr ea 10.0 at Siwa e 11.2 twar Pl DhokP 12 cen Po Rohtas Mio at ha Chinji n Tatrot 14 14.2 i gr Chinji r Na Hasnot 16 we Sethi Lo Kamlial r Ghabir River um Rive nge Jhel 18 Salt Ra 40km 32 o o o 72 73o 74 o 73 18E B 0 1.5 3 N H14 kmkm HH1313 H12 HH1919 H11H11 H18H18 H1H166 H20H20 HH1515 H7 H5H HH88 H4 HH66 o H3 32 49N HH2323 HaHasnot DhokDhok Pathan H2 MiMiddleddl Formation DhokDhok Pathan Upperper Formamation RRecenecent Alluvium H1 LocaliLocalities RiR ver Figure 1. A) A simplified map of the Potwar Plateau in northern Pakistan, with chronostratigraphic context of the Siwaliks Neogene-Quaternary deposits (data from Behrensmeyer & Barry 2005; Dennell 2008; Nanda 2008); reference localities of the Siwaliks are shown with circles. B) A simplified geologic map of the Hasnot area with indicated localities around the village Hasnot (H = Hasnot), from where the studied material was recovered. 309 KHAN et al. / Turkish J Earth Sci Table 1. Mammalian faunas of Hasnot (referred data are taken Table 1. (continued). from Colbert 1935; Pilgrim 1937, 1939; Hussain 1971; Sarwar 1977; Bernor & Hussain 1985; Pickford 1988; Akhtar 1992; Rhinocerotidae Farooq et al. 2007a, 2007b, 2007c, 2007d; Khan et al. 2009; Ghaffar et al. 2010, 2011; Khan 2010). Many species are under Chilotherium blanfordi Chilotherium intermedium taxonomic revision. Subchilotherium intermedium Alicornops sp. Cercopithecidae Brachypotherium perimense Cercopithecus hasnoti Macacus sivalensis Suidae Tetraconodon magnus Propotamochoerus ingens (?) Rodentia Propotamochoerus hysudricus Rhizomys sivalensis Rhizomys sp. Hippopotamodon sivalense Hippopotamodon vagus Hystrix sivalensis Sivahyus punjabiensis Hippohyus lydekkeri Carnivora Anthracotheriidae Amphicyon lydekkeri Indarctos punjabiensis Microbunodon silistrensis Merycopotamus dissimilis Promellivora punjabiensis Enhydriodon falconeri Tragulidae Sivaonyx bathygnathus Vishnuictis salmontanus Dorcabune anthracotherioides Dorcabune nagrii Ictitherium sivalense Hyaenictitherium indicum Dorcatherium majus Dorcatherium minus Lycyaena macrostoma Lycyaena macrostoma-cinayaki Dorcatherium minimus Precrocuta carnifex Precrocuta gigantea Cervidae Precrocuta gigantean-latro Adcrocuta eximia Rucervus simplicidens Cervus triplidens Mellivorodon palaeindicus Acluropsis annectans Cervus sivalensis Cervus punjabiensis Paramachairodus orientalis Felis sp. Cervus rewati Propontosmilus sivalensis Giraffidae Proboscidea Bramatherium megacephalum Bramatherium perimense Dinotherium indicum Paratetralophodon hasnotensis Giraffa punjabiensis Tetralophodon falconeri Tetralophodon punjabiensis Bovidae Zygolophodon chinjiensis Choerolophodon corrugatus Taurotragus latidens Tragoportax salmontanus Anancus perimensis Stegolophodon latidens Tragoportax punjabicus Tragoportax browni Stegolophodon cautleyi Stegodon bombifrons Proleptobos birmanicus Selenoportax vexillarius Stegodon cliftii Stegodon elephantoides Selenoportax lydekkeri Pachyportax latidens Equidae Pachyportax giganteus Gazella lydekkeri Cormohipparion antelopinum Cormohipparion theobaldi Gazella padriensis Elaschistoceras khauristanensis Sivalhippus perimense Hipparion sp. Eotragus sp. ?Tragoportax curvicornis 310 KHAN et al. / Turkish J Earth Sci material of Giraffidae, including B. megacephalum and B. Genus Bramatherium Falconer, 1845 grande. The lithostratigraphy of the area was described in Synonymy: Hydaspitherium Lydekker, 1877 detail by Pilbeam et al. (1977) and Barry et al. (2002). It ? Helladotherium Gaudry, 1860 consists of sandstones with alternate clays and scattered Vishnutherium Lydekker, 1876 conglomerates in the lower part and conglomerates with Type species. Bramatherium perimense Falconer, 1845. sandstones and orange-brown clays in the upper part. Diagnosis. Colbert, 1935: 348. Geographic distribution. Late Miocene to Early Pliocene 2. Materials and methods of the Pakistan-Indian Subcontinent and the Greco- The studied material is stored at the Palaeontology Iranian province (Lydekker 1876; Pilgrim 1910; Matthew Laboratory of the Zoology Department of the University 1929; Colbert 1935; Geraads & Güleç 1999; Sehgal & of Punjab in Lahore, Pakistan, and the Palaeontology Nanda 2002; Bhatti 2004; Khan 2007). Laboratory of the Government College University in Bramatherium megacephalum (Lydekker, 1876) Faisalabad, Pakistan. The material was discovered by a Figure 2; Table 2 team of members from the Palaeontology Laboratories Synonymy: Hydaspitherium megacephalum Lydekker, of the Government College University, Faisalabad, and 1876 the University of Punjab, Lahore, Pakistan, during the Vishnutherium iravaticum Lydekker, 1876 past few years from outcrops near Hasnot, northern Hydaspitherium birmanicum Pilgrim, 1910 Pakistan (Figure 1A). The ascribed material comes from Type specimen. GSI