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GIRAFFA (GIRAFFIDAE, MAMMALIA) FROM THE LOWER SIWALIKS OF PAKISTAN.

Byline: K. Aftab, M. A. Khan, M. A. Babar, Z. Ahmad and M. Akhtar

ABSTRACT

New remains of Giraffa priscilla are recorded from the Middle Miocene localities of Pakistan. The material originates from Chinji Rest House, Rakh Wasnal, Dhok Bun Amir Khatoon, Dhulian, Ghungrilla, Dial, Lava, Phadial, Bhelomar and Ratial. These localities are also well known for the rich Middle Miocene mammalian fauna of the Siwaliks. The estimated age of these localities is 14.2-11.2 Ma, belonging to the Chinji Formation of the Lower Siwaliks. The findings contribute to our understanding of the presence of this species in the Middle Miocene of Pakistan.

Key words: Mammalia, Vertebrates, Giraffa priscilla, Miocene, Siwaliks.

INTRODUCTION

Siwalik Hills are famous for its mammalian fossil fauna (Pilgrim, 1910, 1911, 1913; Matthew, 1929; Colbert, 1935; Sarwar, 1977; Thomas, 1984; Akhtar, 1992; Barry et al., 2002; Bhatti et al., 2007, 2012a, b; Khan et al., 2008, 2009, 2010, 2011, 2012, 2013). The fossils of Giraffidae are reported throughout the Siwalik Early Miocene - Late Pleistocene. The Siwalik giraffids may be placed in three subfamilies i.e. Paleotraginae, Sivatheriinae and Giraffinae. Paleotraginae comprises the genus Giraffokeryx. Sivatheriinae includes the genera Sivatherium, Bramatherium, Helladotherium and Hydaspitherium. Giraffa is placed in the subfamily Giraffinae. These three subfamilies emerged simultaneously but their migration to the Siwalik region occurred at different times. Palaeotragines and Giraffines came earlier than the Sivatheriines (Akhtar et al., 1991).

Different species of Giraffa have been reported from Ethiopia, Kenya and Tanzania during Neogene and Quaternary (Geraads, 1987, 1988; Geraads et al., 2004a, b). Giraffa sivalensis was a descendent of Giraffa punjabiensis and had an Early Pleistocene age (Churcher, 1978; Solounias, 2007). It was extinct from the Upper Siwaliks of Pakistan by the Middle Pleistocene (Mitchell and Skinner, 2003). In Africa, Giraffa first appeared in Early Pliocene (Geraads et al., 2013). Fossils of Giraffa camelopardalis were recovered from Ethiopia (Asfaw et al., 2002), Israel (Haas, 1966; Bar-Yosef and Tchernov, 1972; Geraads, 1986), Northern Africa (Romer, 1928; Howe and Movius, 1947; Arambourg, 1952, 1979; Singer and Bone, 1960; Geraads, 1981), Central Africa (Brunet and M.P.F.T, 2000), Eastern Africa (Kent, 1942a, b; Vaufrey, 1947; Cooke, 1963) and Western Africa (Joleaud, 1936).

Giraffa jumae identified from Turkey, Africa and Middle East (Leakey 1965; Harris et al., 1988; Bruent et al., 1998; Geraads, 1998; Suwa et al., 2003; Wynn et al., 2006). Giraffa pygmaea was smaller than G. camelopardalis and G. stillei. It has been recognized from Ethiopia, Kenya and Malawi (Taieb et al., 1976; Kalb et al., 1982; Harris, 1991; Schrenk et al., 1993; Bromage et al., 1995). Giraffa stillei was larger than G. jumae and G. camelopardalis. It is known from Ethiopia and Africa (Dietrich, 1942; Geraads, 1994; Ward et al., 1999). Arambourg (1947) diagnosed that it was synonym for G. gracillis.

Geography and geology: The Siwalik formations can be differentiated by percentage of sand. In the Chinji Formation, sand is less than 50%. In this Formation cross-bedded facies, sandstone facies, inter-bedded mudstone facies, siltstone and cross-laminated facies have been identified (Kafayat Ullah, 2009). Calcareous nodules are also present which vary in size (Behrensmeyer and Tauxe, 1982; Badgley, 1986). The fossils are collected from the ten Middle Miocene localiteis of the Chinji Formation; a detailed geography and geology of these localities have been reported in the earlier articles (e.g. Sarwar (1973, 1977, 1990), Barry et al. (2002), Cheema (2003), Khan et al. (2005, 2008, 2011, 2012).

MATERIALS AND METHODS

Giraffa priscilla was the oldest known Middle Miocene Lower Siwaliks giraffid (Matthew, 1929; Harris, 1991; Basu, 2004; Bhatti et al., 2012b). New remains of Giraffa priscilla have been reccorded from ten localities i.e. Chinji Rest House, Rakh Wasnal, Dhok Bun Amir Khatoon, Dhulian, Ghungrilla, Dial, Lava, Phadial, Bhelomar and Ratial of the Lower Siwaliks of Pakistan (Fig. 1). The sample provides new evidence of normal variability of giraffid teeth, may in resulting from a mixing of taxa, a previous hypothesis (Geraads, 1989).

The terminology and identification follow Hamilton, (1973), Janis and Scott (1987), Gentry (1994) and Khan et al. (2009).

SYSTEMATIC PALAEONTOLOGY

Superfamily GIRAFFOIDEA Gray, 1821

Family GIRAFFIDAE Gray, 1821

Subfamily GIRAFFINAE Zittel, 1893

Genus GIRAFFA Brisson, 1756

Geographic distribution. Giraffa is best known from Siwaliks of Pakistan and India, Ethiopia, Kenya, Malawi, Tanzania, South Africa and Uganda (Pilgrim, 1910; Hopwood, 1934; Colbert, 1935; Dietrich, 1942; Arambourg, 1947; Cooke, 1963; Leakey, 1965; Hendey, 1968, 1969; Mawby, 1970; Taieb et al., 1976; Pickford, 1986, Harris et al., 1988; Harris, 1991; Schrenk et al., 1993; Geraads, 1994; Bromage et al., 1995; Ward et al., 1999; Basu, 2004; Bhatti et al., 2007, 2012b; Khan et al., 2005, 2010).

Giraffa priscilla Matthew, 1929

Stratigraphic range. Lower Siwaliks (Matthew, 1929; Colbert, 1935; Basu, 2004; Bhatti, 2005; Khan et al., 2012).

New material (in parenthesis the inventory number and the locality name are given): Upper dentition: rP2 (GCUPC 1155/09, Phadial), rM1s (GCUPC 1174/09, Dial; GCUPC 1157/12, Bhelomar), lM2s (GCUPC 1142/09, Ratial; GCUPC 1159/12, Dhulian; GCUPC 724/12, Dial; GCUPC 1154/12, Lava; GCUPC 1189/12, Ghungrilla), rM2s (GCUPC 1139/12, Dial; GCUPC 730/09, Dhulian; GCUPC 906/07, Lava; GCUPC 1186/12, Dhok Bun Amir Khatoon), lM3(GCUPC 491/02, Chinji Rest House; GCUPC 1121/12, Phadial; GCUPC 490/02, Wasnal); Lower dentition: lI3 (GCUPC 1151/12, Dhok Bun Amir Khatoon), lM2 (GCUPC 673/09, Ratial), rM2 (GCUPC 1168/12, Phadial), lM3 (GCUPC 729/05, Wasnal).

Description

Upper dentition: The P2 is rectangular [Fig. 2(1)]. The enamel layer is rugose and somewhat shiny. The parastyle and the rib is strong. The fossette is large and somewhat linear. The molars are broad crown (Figs. 2-4). A thick layer of cingulum is present antero-lingually (Figs 2, 3). The postprotocrista is longer than the preprotocrista. The metaconule is present posterior to the protocone. The premetaconule crista is longer than the postmetaconule crista. It appears to be adjoining with protocone at its anterior end. It is supported by a thick layer of cingulum posteriorly. The paracone is contiguous with metacone through a narrow channel at the posterior side. The paracone extends forward and folds backward anteriorly to produce a very small parastyle. The meta-and mesostyles are incipiently developed. The transverse valley is open lingually. The longitudinal valley is linear antero-posteriorly.

Lower dentition: The left lower incisor is in middle wear and spade shape [Fig. 4(3)]. The enamel boarder is pinched antero-posteriorly and shows a pressure mark at the base which indicates that it is a last incisor. Lingually, the enamel is rough at outer side perhaps due to weathering whereas it is crenulated at the inner side. It has large and wide cutting furrows at the posterior edge, which is divided into major and minor lobes. The anterior lobe is narrow whereas the central and posterior ones are wide. The tooth is laterally compressed showing broad pressure marks caused by the adjacent teeth. The anterolabial side of the tooth is simple, convex and oriented backwardly.

The lower molars are rectangular in outline, nicely preserved and moderately worn teeth [Fig. 4(4-6)].

The major conids are not in straight line. The preprotocristid is longer than the postprotocristid. The both cristids of the hypoconid are contiguous with the protoconid and the entoconid antero-posteriorly. The enamel lining of the metaconid is inclined posteriorly. It is backwardly directed to form a vertical pillar like mesostylid. The metaconid is folded at lingual middle side to form a median rib. The metaconid extends anteriorly and entoconid posteriorly and outwardly to form a thick pillar like metastylid. The entoconid is also supported by a thick median rib. The hypoconulid is supported by a stylid in m3.

COMPARISON AND DISCUSSION

The studied specimens are brachydont and small in size having thick enamel sculpture, so they resemble best with the Lower Siwalik genera Giraffokeryx or Giraffa (Colbert, 1935; Bhatti, 2005; Khan et al., 2010). Giraffokeryx have weakly developed styles and median ribs; the stylids are absent; the spur can be observed in the anterior fossette (Fig. 5). The major cusps and conids are in a straight line (Pilgrim, 1910; Matthew, 1929; Colbert, 1935; Bhatti, 2005; Bhatti et al., 2012a). The genus Giraffa has strong and pillar like styles, clear stylids, strong median ribs and comparatively broad crown. The spur is absent in the anterior fossette and major cusps and conids are not in a straight line. It differs from other genera of the Lower Siwaliks by having reduced posterior half of the tooth (Pilgrim, 1911; Matthew, 1929; Colbert, 1935; Bhatti, 2005; Bhatti et al., 2012b).

The studied material has strong styles/stylids and median ribs, broad crown and obliquely present cusps/conids (Fig. 2-5, Table 1). The posterior half of the tooth is also reduced as compared to Giraffokeryx. These features resemble with Giraffa priscilla (Pilgrim, 1911; Matthew, 1929; Colbert, 1935; Basu, 2004; Bhatti, 2005; Bhatti et al., 2012b) Giraffa priscilla is only recorded from the Lower Siwaliks of Pakistan. However a few specimens have been reported from the base of the Nagri Formation (Middle Siwaliks). The species completely disappeared around 10 Ma from the Subcontinent (Khan et al., 2012).

Table 1. Comparative dental measurements (mm) of Giraffa priscilla. *the studied specimens. Referred data are taken from Pilgrim (1911), Matthew (1929), Colbert (1935), Bhatti (2005), Bhatti et al. (2012b) and Khan et al. (2012).

Taxa###Number###Nature###Length###Width###W/L

###(mm)###(mm)

G. priscilla###GCUPC 1155/09*###P2###22.2###14.9###0.67

###GCUPC 1174/09*###M1###24.0###24.0###1.00

###GCUPC 1157/12*###M1###26.0###26.0###1.00

###GCUPC 1142/09*###M2###25.6###28.9###1.13

###GCUPC 1159/12*###M2###28.6###29.0###1.01

###GCUPC 724/12*###M2###25.0###28.1###1.12

###GCUPC 1154/12*###M2###26.4###29.6###1.12

###GCUPC 1189/12*###M2###26.3###29.2###1.11

###GCUPC 1139/12*###M2###25.9###28.7###1.11

###GCUPC 730/09*###M2###26.2###29.1###1.11

###GCUPC 906/07*###M2###26.0###28.8###1.11

###GCUPC 1186/12*###M2###25.2###28.1###1.12

###GCUPC 491/02*###M3###28.8###30.1###1.05

###GCUPC 1121/12*###M3###28.5###29.1###1.02

###GCUPC 490/02*###M3###31.2###31.9###1.02

###GCUPC 1151/12*###I3###13.6###6.7###0.49

###GCUPC 673/09*###M2###26.8###16.6###0.62

###GCUPC 1168/12*###M2###26.0###15.8###0.61

###GCUPC 729/05*###M3###41.2###22.5###0.55

###PUPC 02/99###P4###19.5###21.0###1.08

###M1###24.0###24.0###1.00

###M2###25.0###28.0###1.12

###PUPC 07/131###M1###25.0###25.0###1.00

###PUPC 07/89###M1###27.0###27.0###1.00

###PUPC 02/9###M3###40.0###17.0###0.43

G. punjabiensis###GSI###P4###20.5###26.6###1.30

###M1###28.2###30.8###1.09

###M2###31.5###34.5###1.10

###M3###32.0###31.5###0.98

###GSI K 13/349###P4###22.0###20.0###0.91

###M1###30.0###24.0###0.80

###M2###32.0###24.0###0.75

###M3###30.0###22.0###0.73

###GSI B 182###M3###29.0###31.0###1.06

###GSI K 13/348###M3###31.0###33.0###1.06

###PUPC 95/23###P4###20.0###23.0###1.15

###M1###31.0###27.0###0.87

###M2###34.0###28.0###0.82

###M3###31.0###23.0###0.74

###PUPC 86/84###M1###21.0###28.0###1.33

###M2###34.0###27.0###0.79

###M3###31.0###24.0###0.77

###GSI###M2###28.3###32.2###1.14

###M3###43###23.2###0.54

###AMNH 19318###M2###26.0###25.0###0.96

G. sivalensis###GSI###M2###33.0###37.0###1.12

###M3###30.4###35.5###1.17

###PUPC 69/123###P4###18.0###22.5###1.25

###PUPC 68/317###M3###27.0###29.0###1.07

###PUPC 67/484###M3###28.0###28.0###1.00

###GSI###M3###43.5###25.2###0.58

Conclusions: Giraffa priscilla is present Middle to earliest Late Miocene of the Siwaliks and it is common faunal elements in the Middle Miocene of the Siwaliks. It can be differentiated from the other Lower Siwalik giraffid genera by strong and pillar like styles and stylids, prominent median ribs and comparatively broad crown. The species was disappeared before the onset of the Dhok Pathan Formation (ca. 10.2 Ma).

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Publication:Journal of Animal and Plant Sciences
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Date:Jun 30, 2016
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