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Short Communication - NEW FOSSILS OF SUIDAE (MAMMALIA) FROM DHOK PATHAN FORMATION OF SIWALIKS, PUNJAB, PAKISTAN.

Byline: F. Y. Dar, K. Aftab, M. A. Babar, M. A. Khan, S. G. Abbas, R. Shahid and M. Asim

Keywords: Artiodactyla, Suiformes, Suidae, Miocene, Siwaliks.

INTRODUCTION

Suids have been classified as even toed ungulates and are abundant in the Siwaliks of Indian Subcontinent. These are diverse in number and species and have been studied by the various researchers from the beginning of paleontological studies in this area (Falconer, 1868; Lydekker, 1883; Pilgrim, 1926; Colbert, 1935; Pickford, 1988; Made, 1996, 1999; Batool et al., 2015). In the past, this family was widespread both in Africa and Eurasia with several genera (Kostopoulos and Sen, 2016., Pickford and Obada, 2016., Spassov et al., 2018., Mors et al., 2019). Some of the species are well-represented in these studies based on their abundant fossils and others have been represented by only a few specimens. Every new specimen of these species is extremely important and valued much because of the information's it provides.

Such a rare species is Tetraconodon magnus, a large sized tetraconodont species, found from the late Miocene of the Siwaliks (Pilgrim, 1926; Pickford, 1988; Made, 1999; Khan et al., 2013). Only 15 specimens are found and some of these are un-described (Made, 1999; Khan et al., 2013). Other species like Hippopotamodon sivalense and Propotamochoerus hysudricus are well-documented with handful fossils record. However, Hippopotamodon sivalense has a close affinity with the European species Microstonyx major and there remains a whether later species arrived in the Siwaliks or both can be synonymized or not (Made and Hussain, 1989., Pickford, 2015). The samples described in this article include the Tetraconodon cf. magnus, Hippopotamodon cf. sivalense and Propotamochoerus hysudricus.

Among these the newly recovered specimen of T. cf. magnus adds the new insights in the morphology of the tooth and jaw of this species and that of Hippopotamodon cf. sivalense confirms the point of view of Pickford (1988) that only a single species is present in the Siwaliks contrary to Made and Hussain (1989). Hence, the new addition in the Siwalik suids is the objective of the article. The present specimens are recovered from the Late Miocene type locality Dhok Pathan (Lat. 33Adeg 07'N: Long. 72Adeg 14'E), Chakwal, Pakistan (Fig. 1). The thickness of these deposits varies from 950 - 1200 m (Barry et al., 2002). The type locality comprises alternate sandstone, claystone/siltstone and rarely with conglomerates (Shah, 1977, 1980; Barry et al., 2002; Bhatti et al., 2012a, b). Magnetic polarity and stratigraphic dating indicate that the age of Dhok Pathan Formation is between 10.1-ca.3.5 Ma.

The lower part of the Dhok Pathan Formation is dated between 10.1-9.0 Ma and the upper part is dated at ca. 9.0-5.5 Ma (Cande and Kent, 1995; Barry et al., 2002).

MATERIALS AND METHODS

Specimens being described in article have been collected from the Dhok Pathan Formation's type locality (Fig. 1). From the gross collection i.e. all the collected specimens, we have selected the suid fossils for this study and removed the sediments with fine needles and brushes. Some of the specimens were partly broken (like 13/370 and 13/63), these were carefully glued with the adhesives after washing them. The specimens were catalogued (e.g. PUPC 16/117) and are kept in Dr. Abu Bakr Fossil Display and Research Centre, Zoology Department, University of the Punjab, Lahore, Pakistan. The catalogue numerator symbolizes the collection year and denominator denotes serial number of that year. Digital Vernier caliper was used for measuring the specimens. The terminology and measurements followed Pickford (1988).

Comparisons were made with specimens present in American Museum of Natural History (AMNH), British Museum of Natural History London (BMNH), the Geological Survey of India (GSI) and the Geological Survey of Pakistan (GSP).

SYSTEMATIC PALAEONTOLOGY

Order ARTIODACTYLA Owen, 1848

Family SUIDAE Gray, 1821

Sub-family TETRACONODONTINAE Lydekker, 1876

Genus TETRACONODON Falconer, 1868

Tetraconodon cf. magnus (Falconer, 1868)

New material: PUPC 13/37, left mandible fragment with a lower third molar (m3).

Description and comparison: PUPC 13/37 is an elongated molar, narrower distally than mesially (Fig. 2(1)). All four cuspids are well differentiated. The protoconid, metaconid, hypoconid and entoconid are organized into two distinct lobes with rounded corners separated from each other by a well-developed median valley. The furrows are indistinct. The protoconid is larger than other cuspids. The third lobe or talonid is present. It is relatively simple and has a length of 26 mm. The pentaconid is very distinct in the molar. The protoendoconulid, hypopreconulid and pentapreconulid are well developed. The tooth shows mesodonty and its enamel is very thick, with visible scratches. The molar has inflated conids with hypopreconulid and pentapreconulid. These features can be recognized in the generous sized Siwalik suid Tetraconodon. Two species were erected by the Siwalik: T. magnus and T. minor (Pickford, 1988; Made, 1999).

Although, morphometrically (Tab. 1), the specimen resembles Tetraconodon magnus, however, being a single specimen collection and is partially broken it has been assigned to T. cf. magnus (Pickford, 1988; Made, 1999; Khan et al., 2013).

Subfamily SUINAE Zittel, 1893

Genus HIPPOPOTAMODON Lydekker, 1877

Hippopotamodon cf. sivalense Lydekker, 1877

New material: PUPC 11/40, right upper first incisor (I1); PUPC 13/374, left upper second premolar (P2).

Description and comparison: PUPC 11/40 is a large spade like tooth with raised labiolingual borders, surrounding a central lingual fossa (Fig.2(2)). A small central rib is not visible at the tip of the central lingual fossa because of wearing. The mesiodistal borders are raised lingually. The occlusal surface of the tooth is beaded and there are light bifurcations near the mesial edge. The parapreconule is clearly visible at the anterior side of the paracone. The basal fossa is present at the base of the protocone. At the mesiolingual border of the tooth a crest is present i.e. preanticlin and next to it there is trough i.e. presyncline. Similarly, at the distal lingual border of the tooth there is a crest i.e. endoanticline and next to it is a trough which is endosyncline. Metapostcrista is clearly visible. At the lingual side of the tooth, two pillar-like ridges prestyle at mesial end and poststyle at distal end are present.

The anteroposterior diameter (DAP) of the tooth is 25.7 mm and transverse diameter (DT) is 10.7 mm. The premolar PUPC 13/374 is an elongated tooth with inflated main cusp (protocone) (Fig.2(3)). The premolar is bordered by a complete cingulum lingually, labial cingulum is lightly formed. The posterolingual cusplet is prominent. The paracone is present mesially. A large fossa is visible on the entire occlusal surface. The parastyle and cingulum is present anteriorly. There is a contact depression posteriorly. The new studied specimens are morphometrically (Tab. 1; Fig. 2(2,3)) associated with Hippopotamodon cf. sivalense.

Genus PROPOTAMOCHOERUS Pilgrim, 1925

Propotamochoerus hysudricus (Stehlin) 1899-1900

New material: PUPC 13/63, right mandible fragment with third premolar (p3) and partial fourth premolar (p4); PUPC 09/95, left mandible fragment with fourth premolar (p4).

Description and comparison: The paraconid is present mesially and entoconid distally in the third premolar (Fig. 2(4)). The protoconid and hypoconid are visible labially while the metaconid is prominent lingually. The anteroposterior cusplets are absent. The 3rd cusp is found distinctively. The parastylid, protostylid, metastylid and entostylid are clearly visible with their respective conids. The distal edge of the premolar is slightly broken lingually. The antero-posterior grooves are present labiolingually.

The p4 is more moralized tooth than the p3 (Fig. 2(5)). The central cusp has migrated lingually to form an inner cusp. The anteroposterior accessory cusps are present. The posterior accessory cusp is enlarged having a border of inflated cingulum. The anterior cingulum, as well as accessory cusp, are moderately high, forming an ac-1 cusp. The p4 is attached with a small portion of broken m1 posteriorly. The socket of a lower canine is also visible. A short, curved symphysis is visible at buccal side of mandibular fragment. A broken incisor is also present at the tip of the symphysis. A mandibular pit can be seen labially. Morphometrically (Tab. 1; Fig. 2 (4,5)), the sample belongs to Propotamochoerus hysudricus.

Suoid indet: New Material: PUPC 13/62, right mandible fragment with roots of second and third molars (m2and m3).

Description and comparison: The molars are in late wear, disappearing the crown (Fig. 2(6)). The length of the fragment is 49.7 mm. Morphologically, the molar features are visible. Therefore, its taxonomy cannot be determined.

Table. 1. Comparative measurements (in mm) of the teeth of Tetraconodon magnus,Hippopotamodonsivalenseand Propotamochoerushysudricus. *the studied specimens. Referred data are taken from Pickford (1988).

Taxa###Inventory No.###Position/Nature###Width###Length###W/L

Tetraconodon cf. magnus###PUPC 13/37*###lm3###29###55###0.53

###B 71###m3###32.5###50###1.54

###B 828###m3###31.4###48###0.65

Hippopotamodon cf. sivalense###PUPC 11/40*###rI1###10.7###25.7###2.4

###GSP 12508###I1###11.3###19.2###1.7

###GSP 330###I1###12.3###-###-

###K 14/394###I1###18###23###1.28

###K 13/328###I1###15###25###1.67

###K 13/345###I1###19###19###0

###K 15/660###I1###17###22###1.3

###PUPC 13/374*###lP2###13###20###1.54

###K 23/105###P2###11.4###18.1###1.59

###B 396###P2###13.3###19###1.43

###K 12/870###P2###12.3###20###1.63

###B 354###P2###11.5###15.6###1.36

###K 42/172###P2###13.4###20.4###1.52

###B 737###P2###13.6###22.1###1.63

###GSP 3789###P2###12.3###-###-

Propotamochoerus hysudricus###PUPC 13/63*###rp3###10.7###15###1.4

###B 32###p3###10###16.7###1.67

###B 39###p3###8###14.7###1.84

###GSP 2807###p3###9.1###15###1.65

###GSP 11598###p3###7.6###-###-

###GSP 457###p3###8.6###15###1.74

###GSP 2503###p3###8.2###15.1###1.84

###GSP 10998###p3###8.2###15.7###1.91

###PUPC 09/95*###lp4###11.7###15.2###1.3

###GSP12343###p4###9.2###16.7###1.82

###GSP 5823###P4###11.3###14.4###1.27

###GSP 9355###p4###11.4###16.6###1.46

###GSP 12732###p4###9.6###16###1.67

###GSP 457###p4###11.4###15###1.32

###GSP 5230###p4###10.8###16.4###1.52

###GSP 48###p4###10.5###-###-

###GSP 2807###p4###10###14.5###1.45

###GSP 7017###p4###12.6###15.6###1.24

###GSP 2503###p4###11.6###13.6###1.17

###GSP 10998###p4###11.6###15.5###1.34

###B 742###p4###10.7###15.3###1.43

###B 39###p4###9.9###13.9###1.4

###B 32###p4###12###18###1.5

###B 715###p4###12.5###17.5###1.4

DISCUSSION

The Siwalik suids form an important assemblage whose ties with the other regions of the world are prominent. Pilgrim (1926) was the first one who discovered vast divergence in the Siwalik Suidae. The Siwalik suids are represented by some endemic genera like Lophochoerus, Tetraconodon, Hippopotamodon, Sivahyus and Hippohyus, whereas some genera are common in Siwaliks and Europe and other parts of the world like Bunolistriodon, Listriodon, Propotamochoerus, Conohyus and Sus (Pilgrim 1926, Colbert, 1935; Pickford, 1988, Made, 1996). The Siwalik suids are also important in that a few genera which evolved in subcontinent, but later migrated to Europe, Africa and other parts of the world like Conohyus, Propotamochoerus, Sivachoerus and Sus (Matthew, 1929; Colbert, 1935; Pickford, 1988; Made 1999).

The African suids have been evolved from the primitive Siwalik Sus (Pickford, 1988). Genus Tetraconodon is represented by three species T. magnus, T. intermedius, and T. minor (Pilgrim, 1926; Pickford, 1988; Made, 1999). All the species are poorly documented and have been based on the isolated dentition mostly. T. minor is considered as the smallest and T. magnus has been regarded as the largest (Pickford, 1988; Made, 1999). The status of the two newly erected species T. malensis by (Thaung-Htike et al., 2005) is yet uncertain. It may represent a variant Conohyus indicus. Pilgrim (1926) and Lydekker (1884) described several specimens of Hippopotamodon sivalense from theLate Miocene of the Siwalik Hills but with different names. Older and younger specimens of this genus was present from about 10 to 7 myr. Hippopotamodon sivalense range is very similar to that of Propotamochoerus hysudricus and apparently Microstonyx.

Nevertheless, it is Late Miocene species which disappearing during Pliocene (Pickford, 1988). According to controlled collection of Yale/Pakistan Expedition, P. hysudricus existing in the Nagri and Dhok Pathan formations along with Sivahyus, Hippohyus and Hippopotamodon sivalense, having age about 10 to 6 myr. Another species Korynochoerus palaeochoerus has been discovered from Europe in the same range (Pickford, 1988).

Conclusions: The Dhok Pathan type locality displays the three contemporaneous and sympatric genera Tetraconodon, Propotamochoerus and Hippopotamodon. Tetraconodon magnus, Hippopotamodon sivalense and Propotamochoerus hysudricus are Late Miocene species that disappeared at the end of Pliocene. The type locality of the Dhok Pathan Formation provides evidence of diversifying suids in the Late Miocene of the Siwaliks.

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Publication:Journal of Animal and Plant Sciences
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Date:Aug 27, 2019
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