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Dorcatherium cf. nagrii from the Chinji Type Locality (Chakwal, Northern Pakistan) of the Chinji Formation, Lower Siwaliks, Pakistan.

Byline: Muhammad Akbar Khan and Muhammad Akhtar

Abstract

This paper reports the first discovery of complete lower molar series ascribed to Dorcatherium cf. nagrii from the Chinji type locality of the Chinji Formation. The material, discovered by the team of Palaeontology of the Punjab University, Lahore, Pakistan in the past year comes from the Middle Miocene continental deposits of the Lower Siwaliks (Chinji Formation, northern Pakistan) dated approximately from 14.2 to 11.2 My, and provides new information about the lower dentition of D. nagrii. The new material extends the presence of D. nagrii to the type locality of the Chinji Formation and suggests a humid habitat with abundant cover, based on D. nagrii and the already recorded taxa from the locality.

Keywords: Siwaliks, tragulids, Chinji Formation, Dorcatherium, Miocene.

INTRODUCTION

The majority of fossil tragulids and the totality of fossil Siwalik species have been included in the extinct genus Dorcatherium, which was first described from the late Miocene of Europe with the type species D. naui (Kaup and Scholl, 1834; Lydekker, 1876; Colbert, 1935; West, 1980; Farooq et al., 2007a,b, 2008). As noted by earlier researchers, this genus embraces the major part of the Miocene-Pliocene diversity of the Tragulidae, with 22 recognized species, exceeding other genera such as Siamotragulus, Dorcabune, Afrotragulus and Yunnanotherium by far in species diversity (ROssner, 2007; Quiralte et al., 2008; Sanchez et al., 2010). Dorcatherium includes a varied array of bunoselenodont and selenodont tragulids that show a great degree of variation in both body size and dental morphology (ROssner, 2007).

Mostly, the variation in the body size has been classically overused to diagnose and assign Dorcatherium species (Arambourg and Piveteau, 1929; Colbert, 1935; Whitworth, 1958; West, 1980; Gaur, 1992; Pickford, 2001; Morales et al., 2003; Farooq et al., 2007a-b, 2008; Quiralte et al., 2008) with much less emphasis put on the description of morphologically diagnostic characters (Morales et al., 2003; Geraads et al., 2005; Hillenbrand et al., 2009). Dorcatherium has a widespread biogeographic distribution that covers Africa, Eurasia, the Greco-Iranian province and the Siwaliks from the early Miocene to the early Pliocene (Arambourg, 1933; Whitworth, 1958; Janis, 1984; Fahlbusch, 1985; Gaur, 1992; Gentry et al., 1999; Pickford, 2001; Morales et al., 2003; Pickford et al., 2004; Farooq et al., 2007a-b, 2008; Quiralte et al., 2008; ROssner, 2007, 2010).

The association of Dorcatherium with Dorcabune, Tragoportax Miotragocerus, Pachyportax, Selenoportax, Giraffokeryx and Hydaspitherium is common in the Siwaliks, implying that they probably had same feeding resources at this time period (Colbert, 1935; Pilgrim, 1937, 1939; Farooq et al., 2007a-d, 2008; Khan et al., 2009, 2010). Bouvrain (1994) suggested that 'Tragoportax' lived in a more wooded environment, because it is more often associated with cervids and tragulids (Farooq, 2006; Khan, 2007; Khan et al., 2010). Merceron et al. (2006) suggests that Tragoportax were engaged in both browsing and grazing. The taxonomic composition indicates the paleoecological conditions of the tragulids compatible with earlier reconstructions of the riverine environment and a covering of woods, bushes and shrubs (Barry et al., 2002).

The Siwalik Dorcatherium is represented by 4 species namely D. minimus, D. nagrii, D. minus and D. majus (Lydekker, 1876; Colbert, 1935;

Prasad, 1968; West, 1980; Gaur et al., 1983). During the past decades, many new Dorcatherium specimens (D. minus and D. majus) have been recovered from the Siwaliks (Farooq, 2006; Khan, 2007; Farooq et al., 2007a-b, 2008), but only two additional species D. minimus and D. nagrii have been recognized (West, 1980; Gaur et al., 1983). Dorcatherium minimus is extremely rare, only two specimens having been collected by West (1980) at Dued Khel (Mianwali district) from the Chinji Formation of the Lower Siwaliks. West (1980) reveals that the only material available to him was an isolated upper third molar and a left astragalus, and he mentions the second upper third molar in addendum. Remains of D. nagrii are normally scarce and they constitute a very rare faunal element in the Siwaliks of Pakistan (Colbert, 1935; Farooq, 2006).

Earlier scientists who have worked on the Siwalik D. nagrii are Colbert (1935), Prasad (1968), Shani et al. (1980), Gaur et al. (1983) and Gaur (1992). Colbert (1935, p. 312) placed the speimens AMNH 19306, AMNH 19613, AMNH 29854 in D. sp. Later on Gaur et al. (1983) erected D. nagrii, based on Colbert's specimens and the specimens collected from Uttar Pradesh and Himachal Pradesh, Indian Siwaliks. Since the discovery of D. nagrii by Colbert in 1935 in northern Pakistan, no one described any material of D. nagrii from the Pakistani Siwaliks. Recently, a mandibular ramus ascribed to D. cf. nagrii has been recovered from the type area of the Chinji Formation. This contribution documents a new lower molar series (m1-3) of rare tragulid D. nagrii recovered from the outcrops south of the Chinji village (Fig. 1).

Biostratigraphy and age

The occurrence of tragulid D. nagrii in the Chinji type locality outcrops, northern Pakistan could indicate the first appearance in the Chinji Formation of the Lower Siwaliks. Earlier specimens of D. nagrii are excavated from the Middle Siwaliks at Nagri level (Colbert, 1935; Gaur et al., 1983; Gaur, 1992). Colbert (1935) attributes the D. nagrii's samples to the localities 5 and 12 miles east of the Chinji Bungalow, and he estimated that they were situated in the upper Chinji Formation. If these were the exact localities, the samples would have an

Eocene origin. The geological map of Colbert (1935) shows the Chinji - Nagri formation boundary further south than is now recognized. New maps with better formation boundaries within the Siwaliks show the areas south of the Chinji Bungalow, which belong to the topmost part of the Chinji Formation (Raza et al., 1983). This erroneous age assignment was based on incomplete locality information of the earlier recovered samples. The new discovery of D. nagrii from the Chinji type locality outcrops enables us to calibrate the stratigraphic position of this species.

The Chinji outcrops have yielded a rich and diversified vertebrate fauna. The fauna from the Chinji Formation includes at present the following species: Sivapithecus sivalensis, S. indicus, Ramapithecus punjabicus, Rhizomyoides punjabiensis, Hyanailouros bugtiensis, Dissopsalis carnifex, Chalicotherium salinum, Listriodon pentapotamiae, Conohyus chinjiensis, Merycopotamus pusillus, Palaeohypsodontus sp., Dorcatherium majus, D. minus, Dorcabune anthracotherioides, Eotragus sp., Miotragocerus gluten, Kubanotragus sakolovi, Sivoreas eremite, Gazella sp., Giraffokeryx punjabiensis, Giraffa priscilla (Matthew, 1929; Colbert, 1933, 1935; Pilgrim, 1937, 1939; Raza, 1983; Akhtar 1992; Barry et al., 2002; Badgley et al., 2008; Khan et al., 2008, 2009, 2010). This faunal association contains enough significant elements to allow comparison with some other middle Miocene to early late Miocene faunas from Europe and Greco-Iranian Province.

The Chinji Formation fauna is clearly of middle Miocene to early late Miocene age, as above-mentioned taxa represent an association of the late middle Miocene and the earliest late Miocene.

MATERIALS AND METHODS

The mandible fragment that is described in this paper has been recovered from the outcrops south of the Chinji village (72deg 22' E, 32deg 41' N), district Chakwal, northern Pakistan (Fig. 1). The outcrops belong to the Chinji Formation of the Lower Siwaliks (Barry et al., 2002; Khan et al., 2008, 2009). The outcrops dominantly comprise bright red and brown orange siltstones interbedded with soft, ash grey sandstones. The sediments were probably deposited in a fluvial environment leaving behind the fine-grained and fossil-bearing flood plains. For a detailed description of the geological and biostratigraphical setting of the area see Behrensmeyer (1987, 1988), Willis (1993), Behrensmeyer et al. (1995) and Barry et al. (2002).

The morphological and metrical characters of the sample are described and their systematic determination is discussed. The specimen is housed in the Zoology Department of the Government College University, Faisalabad, Pakistan. The specimen is registered by the year and a serial catalogued number (e.g. PC-GCUF 10/23). All measurements are given in millimeters (mm). The dental length (L) and width (W) were measured at the occlusal level. The tooth height (H) equals the height of the metaconid on lower molars. The terminology and measurement of the tragulid teeth follow Gentry et al. (1999) and ROssner (2010).

Abbreviations

PC-GCUF, Paleontological Collection of

Government College University Faisalabad; PUPC, Punjab University Paleontological Collection; AMNH, American Museum of Natural History; My, million years; ma, million years ago; m, lower molar; L, largest length; W, largest width; H, maximum height; L(hypc), length of hypoconulid; W(hypc), width of hypoconulid; H(hypc), height of hypoconulid; r, right; l, left.

SYSTEMATIC PALAEONTOLOGY

Suborder Ruminantia Scopoli, 1777

Infraorder Tragulina Flower, 1883

Family Tragulidae Milne-Edwards, 1864

Genus DORCATHERIUM Kaup and Scholl, 1834

Dorcatherium cf. nagrii Gaur, Vasishat, Suneja and Chopra, 1983 (Fig. 2, Table I)

Material

PC-GCUF 10/23 a right mandible fragment with m1-3 (m1: L = 8.0, W = 4.8, H = 5.0; m2: L =

Table I: Comparative measurements of the cheek teeth of the Siwalik Dorcatherium in mm (millimeters).###The studied

###specimens. Reference data are taken from Colbert (1935), Gaur et at (1983) and Farooq et at (2007a-b, 2008).

###Number###Nature/Position###Length###Width###WIL ratio

D. cf. nagrii###PC-GCUF 10/23###8.00###4.8###0.60

###rnI2###8.4###5.0###0.59

###rm3###12.6###5.0###0.39

D. nagrii###AMNH 19306###rm2###8.00###5.0###0.62

###rm3###11.5###5.0###0.43

###GSI 18079###ml###6.50###3.0###0.46

###m2###6.6###3.0###0.45

D. minus###PUPC 02/158###rml###10.6###6.70###0.63

###GSI B594###rml###10.8###6.80###0.62

###PUPC 68/294###rm2###11.0###6.40###0.58

###PUPC 68/311###rm2###10.0###6.60###0.60

###PUPC 68/3 12###1m2###10.0###6.20###0.62

###PUPC 68/3 13###rm2###10.2###6.70###0.65

###PUPC 02/158###rm2###12.7###8.20###0.64

###AMNH 19365###rm2###13.0###7.50###0.57

###AMNH 19366###rm2###12.0###7.50###0.62

###GSI B594###rm2###12.5###7.50###0.60

###PUPC 68/294###rm3###16.1###6.80###0.42

###PUPC 68/311###m3###14.8###7.80###0.53

###PUPC 68/3 13###1m3###15.6###7.40###0.47

###PUPC 83/610###1m3###18.5###8.50###0.45

###PUPC 83/626###1m3###12.5###8.00###0.64

###PUPC 84/82###rm3###18.4###8.30###0.45

###PUPC 85/35###1m3###15.0###7.00###0.64

###PUPC 85/59###1m3###14.2###7.00###0.49

###PUPC 86/266###rm3###14.5###6.40###0.44

###PUPC 96/66###1m3###13.0###6.30###0.48

###PUPC 02/158###rm3###18.5###8.70###0.46

###AMNH 19365###rm3###18.0###8.00###0.44

###AMNH 19366###rm3###16.0###8.00###0.50

###GSI B594###rm3###16.7###8.30###0.49

D. majus###PUPC 86/02###ml###14.3###9.00###0.62

###PUPC 86/05###ml###13.0###9.30###0.71

###AMNH 19524###ml###13.5###9.00###0.66

###GSI B593###ml###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

###PUPC86/05###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

###PUPC 84/115###1m3###24.0###11.0###0.45

###PUPC 86/2###1m3###25.1###11.0###0.43

###PUPC 86/3###1m3###25.0###11.4###0.45

###PUPC 86/152###1m3###23.0###11.0###0.47

###PUPC 96/64###1m3###22.0###11.0###0.50

###PUPC98/61###1m3###16.0###11.0###0.68

###AMNH 19939###1m3###25.5###12.0###0.47

###GSI B593###1m3###25.0###11.4###0.45

8.4, W = 5.0, H = 5.3; m3: L = 12.6, W = 5.0, H = 7.0, L(hypc) = 2.0; W(hypc) = 2.0, H(hypc) = 3.0).

Description

PC-GCUF 10/23 is a dainty broken hemimandible with bunoselenodont morphology (Fig. 2; Table I). The ascending ramus begins right at the posterior end of the hypoconulid, so there is no gap between m3 and the ascending ramus. The hemimandible is shallow and the ascending ramus is broken off. The lingual surface of the jaw is marked by a shallow groove distally. The labial side of the hemimandible is clearly convex anteriorly and the lingual side is slightly concave below m1. The anteroposterior length of the hemimandible is 43 mm and the depth of the hemimandible at m1 is 14 mm. The length of the molar series is 31 mm. The lingual conids are strongly compressed transversely and higher than the labial ones, which are crescent- shaped. m1: The tooth is complete, sub-hypsodont, narrow-crowned and has the typical rounded shape of the anterior contact facet for the premolar (Fig. 2). It is in early stage of wear. All the conids are well preserved.

The lingual conids are slightly more pointed and higher than the labial conids. The anterior cingulid is well developed. The anterior fossette is not crescent shaped but it is somewhat compressed and straight. However, the posterior fossette is crescentic and opens disto-lingually. The vestigial ectostylid is present at the base of the transverse valley. A slight cingulid is present on the base of the labial side. The Dorcatherium fold is present. It is formed by the bifurcation of the post- metacristid resulting formed a 'M'- structure, a diagnostic feature of Dorcatherium (Colbert, 1935; ROssner, 2007, 2010). The lingual ribs are separated by a vertical groove. m2: This tooth is sub-hypsodont, slightly worn and rugose molar (Fig. 2). The conical metaconid is pointed and higher than the protoconid and the hypoconid. The entoconid is broken at the apex. The tubercle-like ectostylid occurs between the labial crescent conids (protoconid and hypoconid). The posterior fossette is crescentic and opens lingually.

The cingulid is present anteriorly. The 'M'-shaped Dorcatherium fold is visible in occlusal view. The anterior rib and the stylid are comparatively weaker than the posterior ones. The pre-metacristid joins the pre-protocristid, while the post-hypocristid contacts the post-entocristid at the base, opening up the posterolingual end of the posterior fossette and forming a weak entostylid. Strong median ribs are present on the lingual side of the tooth and a narrow vertical groove occurs between the two conids. The talonid is slightly larger in length than the trigonid. The stylids are not very bulbous.

m3: The third molar is unworn and completely preserved (Fig. 2). It is higher than m1 and m2. All the conids are pointed and their tips are sharp. No ectostylid occurs in the transverse valley. The enamel is rugose especially on the buccal side of m3. All the crown features are preserved and the 'M' structure of the Dorcatherium fold can be seen pretty clearly. The anterior fossette on the occlusal surface is isolated mesially and the posterior one is comparatively narrow. The fossettes are deeper than those of m1 and m2. The anterior cingulid and the weak metastylid are present anteriorly. The anterior and posterior ribs are strong. The hypoconulid is present, isolated, obliquely situated between entoconid and hypoconid, and comparatively lower in height than the other four major conids. The hypoconulid has a shallow vertical groove mesially. This groove opens lingually. The antero-labial end of the hypoconulid contacts the post-hypocristid at the midline of the tooth.

There is a short, low crest along the lingual edge of the hypoconulid which fails to reach the posterior of the entoconid, leaving the lingual wall incomplete.

Comparison

The semi-selenodonty pattern of the molars confirms the inclusion of this specimen in Tragulidae (Colbert, 1935; Pickford, 2001; Morales et al., 2003; Pickford et al., 2004). The presence of cingulid, stylids and Dorcatherium-fold (M- structure) are characters that correspond to genus Dorcatherium (Lydekker, 1876; Colbert, 1935; Whitworth, 1958; Gaur, 1992; Pickford et al., 2004; Farooq et al., 2007a, b, 2008; ROssner, 2007, 2010). Dorcatherium is represented by four Siwalik species D. minimus, D. nagrii, D. minus and D. majus (Lydekker, 1876; Colbert, 1935; West, 1980; Gaur et al., 1983; Farooq et al., 2007a-b, 2008).

Morphological and metrical features of the specimens clearly indicate a small-sized Miocene tragulid (Table I, Fig. 3). The teeth are too small for D. minus and D. majus (Fig. 3) (Farooq et al., 2007a-b, 2008). Cingulid and stylid cingula are weaker than those of D. minus and D. majus. In addition, the obliquely situated hypoconulid with an incomplete lingual wall can be observed in the small-sized Siwalik Dorcatherium. Nevertheless, Dorcatherium minimus is small enough to exclude the studied sample (West, 1980).

The overall pattern of the described molars is clearly different from that of D. minus and D. majus, and much more similar to the pattern observed in D. nagrii. Morphometrically, the described sample agrees with the earlier specimens of D. nagrii (Table I, Colbert, 1935; Gaur et al., 1983; Gaur, 1992). The teeth positions clearly overlap in size with the already studied material and fall pretty well within the range of D. nagrii (Fig. 3, Table I). Lacking adequate material, it is referred to as D. cf. nagrii. Nevertheless, more material is needed for precise identification.

CONCLUSIONS

The presence of Dorcatherium indicates more or less closed and humid habitats (Kohler, 1993; Gentry, 2005; Eronen and ROssner, 2007). The faunal composition suggests a humid habitat pocket with abundant cover indicating the dominance of forested landscapes during the middle Miocene and the early late Miocene times of the Chinji Formation. The presence of D. nagrii with D. minimus, D. minus and D. majus in the Chinji Formation suggests a strong connection to wet, forested habitats with dense understory, where the animals could hide in vegetation or water from predators (Colbert, 1935; West, 1980; Farooq et al., 2007a-b; Badgley et al., 2008; Khan et al., 2008). A significant representation of tragulids with adaptations to ecotonal wet and swampy habitats indicates humid conditions in the Chinji Formation of the Lower Siwaliks.

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Muhammad Akbar Khan1 and Muhammad Akhtar2, 1Zoology Department, Government College University, Faisalabad, Pakistanm, 2Palaeontology Laboratory, Zoology Department, Quid-e-Azam Campus, Punjab University, Lahore, Pakistan
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Author:Khan, Muhammad Akbar; Akhtar, Muhammad
Publication:Pakistan Journal of Zoology
Article Type:Report
Geographic Code:9PAKI
Date:Dec 31, 2011
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