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New Fossil Locality in the Middle Miocene of Lava from the Chinji Formation of the Lower Siwaliks, Pakistan.

Byline: Muhammad Akbar Khan Muhammad Akhtar, Abdul Majid Khan, Abdul Ghaffar, Mehboob Iqbal and Khizar Samiullah

Abstract. - The middle Miocene locality in the fluvial deposits was discovered in the south-eastern part of Lava village (Chakwal, northern Pakistan). The mammalian assemblage originating from this locality is described. The faunal association consists of rhinocerotids Gaindatherium browni and Brachypotherium fatehjangense, a suid Listriodon pentapotamiae , a caprine Kubanotragus sp., a bovid Helicoportax cf. tragelaphoides, a tragulid Dorcatherium, a giraffid Giraffokeryx punjabiensis and a proboscidean Deinotherium cf. pentapotamiae. The assemblage indicates the locality is middle Miocene in age (ca. 14-11 Ma) and it displays regional characteristics of the Chinji Formation of the Lower Siwaliks, Pakistan.

Key Words: Lava, Middle Miocene, Chinji Formation, Lower Siwaliks, vertebrates.

INTRODUCTION

A fossil locality has been found near Lava village which is 14 km east of Rawalpindi - Mianwali highway. The fossil site is 11 km south east of the village Lava, district Chakwal (Punjab, Pakistan) and east of Dera Rehmatay Aali along the road opposite to Kas Badri from which we obtained fossils in a very dense concentration over a small area (Fig. 1). The fossils are fragmentary mostly postcranial than the cranial ones. The weathering cracks, abrasion marks and bite marks are frequently noted on the specimens. The site is highly fossiliferous and seems to be exposed since long time. Owing to the long exposure fragmentary bones and enamels are present all over the site. Nearly every sandstone has some embedded vertebrate remains. The locality is characterized by sandstone and reddish shale which is characteristic of Chinji Formation of the Siwaliks (Fig. 1). The Formation consists primarily of grey, sandy to muddy fluvial deposits (Barry et al., 2002).

The section consists of shales, siltstones and sandstones. The fluvial sediments display large lateral and vertical variations in their degree of cementation. A few cheek teeth fragments, some undeterminable tooth splinters and some fragmentary postcranial bones are found in the section. This paper presents the study of some new vertebrate fossil material from the discovered locality of the village Lava collected in 2007 - 2009 by the 'team' formed by the Zoology Department of Punjab University, Lahore and the Zoology Department of GC University, Faisalabad, Punjab, Pakistan.

The specimens were catalogued and given a number comprising year of collection and a serial catalogue number ( e.g. 07/104). Various measurements of the studied specimens in millimeters were taken with the help of metric vernier caliper. Tooth length and width were measured at occlusal level. Heights were measured on the mesostyle of the upper molar, the metastylid of the lower molar and the protoconid of the lower premolar. The upper case letter denotes upper dentition and the lower case letter denotes lower dentition. Measurements given for teeth are occlusal length and occlusal width. All the described fossils are housed in the Palaeontological Collection of the Punjab University, Lahore (PUPC) and the Palaeontological Collection of GC University, Faisalabad (PC-GCUF).

Abbreviations

PUPC, Punjab University Paleontological Collection; PC-GCUF, Palaeontological Collection of GC University, Faisalabad; C, canine; d, deciduous; H, height; I, incisor; L, largest length; l, left; M, molar; Ma, million years; MN, Mein Zones; P, premolar; r, right; W, width.

SYSTEMATIC ACCOUNT

Order Perissodactyla Owen, 1848 Family Rhinocerotidae Gray, 1825 Subfamily Rhinocerotinae Gray, 1821 Tribe Rhinocerotini Gray, 1821 Subtribe Rhinocerotina Owen, 1845 Genus GAINDATHERIUM Colbert, 1934

Gaindatherium browni Colbert, 1934

Referred material PUPC 07/101 - rM1; PUPC 07/102 - rM2.

Description PUPC 02/101 is a well preserved first molar and slightly worn (Fig. 4A). PUPC 02/102 is complete, newly erupted and excellently preserved second molar (Fig. 4B). The median valley is filled with matrix in the second molar. The enamel is rugose and the traces of the cement are present all around the molars. The anterior cingulum is well developed and serrated. The posterior cingulum is limited around the postfossette. There is no cingulum at the lingual and buccal face of the tooth. The postfossette is deep and funnel shaped. The median valley is wide open and the protocone and the hypocone are far apart from each other. There is no trace of antecrochet or crista. However a delicate crochet extends into the median valley from the apex of the metaloph. A weakly developed mesostyle is present. The parastyle and the paracone fold are well developed and prominent. The metastyle is also well developed and the metacone rib is also weakly developed.

The protoloph and the metaloph are oriented obliquely to the ectoloph. The concave ectoloph is present behind the paracone fold. A convexity corresponding to the mesostyle is also present. The comparative measurements of all the specimens are provided in Table I.

Table I.- Comparative measurements (cm) of the cheek teeth of G. browni.

G. G. G. G.vidali

Specimen###browni###browni###browni###Heissig###

###present###Heissig###Colbert###(1972)###

###collection###(1972)###(1934)###

PUPC###L###44.5###43.0###40.0###34.0###

07/101 -###W###51.0###53.0###51.0###44.0###

rM1###H###45.0###46.0###X###41.0###

PUPC###L###45.4###46.0###42.0###-###

07/102 -###W###50.5###52.0###52.0###-###

rM2###H###42.0###42.0###-###-###

Discussion

Colbert (1934) described genus Gaindatherium with G. browni from the Chinji Formation of the Lower Siwaliks. Heissig (1972) worked on the Siwalik rhinoceros and described two successive species of Gaindatherium G. browni and G. vidali from the Chinji Formation and from the Nagri Formation of the Siwaliks, respectively. The upper molars show the features of the diagnosis; a simple crochet which is strongly angled against the metaloph, the paracone is narrow and backward, not clearly demarcated, and the parastyle and parastyle folds are strongly developed. Features of the molars, like presence of a stronger anterior cingulum and a protocone fold, presence of a strong metacone rib, are the characteristics of the G. browni. The inner cingulum in G. vidali is weaker than G. browni. The paracone and the metacone are also weakly demarcated. The dimensions of the present material are closer to G. browni and it differs from G. vidali in crown morphology and size (Table I, Fig. 2).

Subtribe Teleoceratina Hay, 1902

Genus BRACHYPOTHERIUM Roger, 1904

Brachypotherium fatehjangense Pilgrim, 1910

Referred material

PUPC 07/104 - ld2, PUPC 07/105 - rP2, PUPC 07/109 - rP2, PUPC 07/106 - lP3.

Description

PUPC 07/104 is broken on the posterior side and fairly identical to the second premolar. It displays a reduced paraconid and open posterior valley. The anterior cingulum is low and asymmetrical. Its height is important on the posterior side and decreases anteriorly (Fig. 4C).

PUPC 07/105 and PUPC 07/109 are partially broken right upper second premolars (Fig. 4D,E).

The premolars are characterized by the open median valley, antecrochet is moderately present, ectoloph is broken and the metaloph is complete. The premolars are in late wear. The cingulum and crista are absent. The crochet is prominently present. The protocone is slightly extended. The posterior fossette is also present.

PUPC 07/106 is in middle wear with broken ectoloph (Fig. 4F). The protocone is slightly extended as compared to the hypocone. The metaloph is incomplete and the protoloph has broken away. The protocone is a strong lingual-anterior pillar, well separated from the hypocone pillar due to the presence of a medisinus. The crista and the crochet are absent. The parastyle has broken off.

Discussion

A reduced paraconid in the second lower premolar is an autapomorphy of Brachypotherium (Antoine, 2002). The morphological features are consistent with B. fatehjangense, known from the early Miocene to the late Miocene in South Asia (Heissig, 1972; Antoine and Welcomme, 2000; Welcomme et al., 2001). An open posterior valley, low and asymmetrical anterior cingulum, the posterior cingulum extends until the posterior groove of the ectolophid are the characteristics correspond to B. fatehjangense. All these characters are observed in the studied material, which clearly identify it to B. fatehjangense. Morpho -metrically, the specimen resembles with specimens originating from the middle Miocene Chinji Formation of the Lower Siwaliks of Pakistan and Myanmar (Table II).

Order Artiodactyla Owen, 1848

Family Suidae Gray, 1821

Subfamily Listriodontinae Simpson, 1945 Genus LISTRIODON von Meyer, 1846

Listriodon pentapotamiae Falconer, 1868

Referred material

PC-GCUF 09/28 - li2; PC-GCUF 09/27 - c; PUPC 07/111 - rp4.

Description

PC -GCUF 09/28 has a stepped cutting edge like spatula (Fig. 4G). The occlusal edge of the tooth is almost straight. The distal edge of the tooth forms a second cutting edge at right angle to the first one. The incisor is mesio-distally compressed posteriorly. There is a prominent centrally placed lingual pillar. The distal edge protrudes slightly forming a shallow scoop like basin between the central pillar and distal edge of the crown.

Table II.- Comparative measurements (cm) of B. fatehjangense. * studied specimens.

Table II.-###Comparative measurements (cm) of B. fatehjangense. * studied specimens.###

Taxon###Specimen###level###Age###Length###Width

B. fatehjangense *###PUPC 07/104 - ld2###Chinji Formation###Middle Miocene###28.1###19.4

B. fatehjangense *###PUPC 07/105 - rP2###Chinji Formation###Middle Miocene###27###29

B. fatehjangense *###PUPC 07/109 - rP2###Chinji Formation###Middle Miocene###26###24

B. fatehjangense *###PUPC 07/106 - lP3###Chinji Formation###Lower Miocene###26###28

B. fatehjangense (Heissig, 1972)###rp2###Chinji Formation###Middle Miocene###27.0###16.0

B. fatehjangense (Chavasseau et al.,2006) rp2###Chaungtha, Myanmar###Middle Miocene###32.3###19.6

PC-GCUF 09/27 is a lower canine with a long permanently growing single root (Fig. 4H). The canine emerges almost horizontally and sweeps outwards and eventually backwards with scrofic cross section. The canine root is long and circular. The enamel is present on the lingual and labial surfaces but it does not extend onto the root. There is a prominent wear facet anteriorly caused by abrasion against the upper canine.

PUPC 07/111 is four cusped lophodont tooth (Fig. 4I). The tooth is molarized with well developed anterior and posterior cingula. The premolar has prominent innenhugel which is closely fused to the main cusp and rectangular occlusal outline. Anteriorly the anterior crested of the main cusped is swollen and descends rapidly towards a well developed but low cingulum. Distally there is a prominent talonid joined lingually and labially by a swollen cingulum. The distal accessory cusplet is two thirds height of the crown. The distal cingulum fades into the labial and lingual surfaces of the crown.

Discussion

The lower incisor shows the typical morphology of listrodonts with stepped edge. The long permanently growing root is present in a male lower canine of L. pentapotamiae (Pickford, 1988).

The listriodonts have a lophodont p4 and well developed anterior and posterior cingula (Pickford and Morales, 2003). The dimensions (Table III; Fig. 3) and the size of the currently documented teeth are close to the teeth of L. pentapotamiae as noted by Pickford (1988). The closely resembling material recovered from the locality is assigned here to L. pentapotamiae.

Table III.- Comparative measurements (cm) of L. pentapotamiae. Referred material is taken from Pickford (1988). * studied specimens.

Lower 2nd###Length###Width###Lower 4th###Length###Width

incisor###premolar###

PC-GCUF###11.3###9.0###*PUPC###15###11

09/28 -i2*###07/111 - p4###

GSP 4527###12.9###9.4###K13/808 -###15.3###12.3

- i2###p4###

K15/537 -###11.4###9.0###K13/436 -###17.4###11.3

i2###p4###

K19/57 - i2###10###11###K23/721-###16.1###12.5

###p4###

K15/587 -###15.4###7###K13/847-###18.4###13

i2###p4###

K41/895 -###13.5###10.5###K14/492-###16.5###11.8

i2###p4###

K15/537 -###12.3###9.7###K16/425-###18.9###14

i2###p4###

K13/795 -###11.7###9.3###Canine###

i2###

K10/407 -###10###6.7###*PC-GCUF###16###13

i2###09/27 - c###

GSP 4461###11.9###9###

- i2###

GSP 4462###12###11.5###

- i2###

GSP 4115###11.5###9.5###

- i2###

GSP 4416###12###10.5###

- i2###

Suborder Ruminantia Scopoli, 1777 Family Tragulidae Milne-Edwards, 1864 Genus DORCATHERIUM Kaup, 1833 Dorcatherium sp.

Referred material

PUPC 07/112 - p4.

Description

The premolar is triangular in lateral view and increase in length from front to back. The tooth is somewhat elongated (Fig. 6A) . The premolar has a strong protoconid linked by a straight longitudinal crest to a small, mesio-lingually centrally located paraconid. The protoconid crest is slightly curved lingually, and joins the more lingually situated paraconid. The hypoconid is situated at the rear of the tooth. The hypoconid is weaker and lower than the protoconid. The posterior side of the hypoconid forming a well marked transverse spur delimits a small, lingually open rounded basin. A very tiny transverse entostylid present at the posterior edge of the crown.

Discussion

The morphology and the orientation of the occlusal outline of the fourth premolar are close to Dorcatherium and differ from those of bovids (Farooq et al., 2008). The size index of the premolar is close to Dorcatherium majus (Table IV). Unfortunately, a mandibular ramus with molars missed on the way from Lava locality to village Lava. The lower molars had well developed Dorcatherium folds. The molars were rectangular in shape and bunoselenodont. Enamel slightly wrinkled. The morphology of the teeth suggests that

they belonged to the Dorcatherium. The lack of material does not allow us to assign species for the remains.

Table IV.- Comparative dental measurements (mm) of fourth lower premolars in Dorcatherium.

Comparative material is taken from Farooq et al. (2008). * studied specimen.###

p4###PUPC###PUPC###AMNH###GSI###* PUPC###

###86/2###86/5###19524###B593###07/112###

L###13.3###13.1###14.5###17.3###16.0###

W###6.0###5.7###5.0###6.2###6.3###

Family Bovidae Gray, 1821

Tribe Boselaphini Knottnerus-Meyer, 1907

Genus HELICOPORTAX Pilgrim, 1937

Helicoportax cf. tragelaphoides

Referred material

PUPC 07/114 - M1; PUPC 09/55 - lp3-m1.

Description

The molar is quadrate. The median ribs are present and the posterior median rib is weaker than the anterior one. The entostyle like basal tubercle is present. The enamel is rugose and has prominent striae (Fig. 6B). The molar has pronounced neck at the base of the crown. The molar has divergent styles and strong buccal folds. The lower molars are narrow crowned and also hypsodont (Fig. 6C). They have pronounced neck at the base like upper dentition. These are in early wear. The p3 is long with separated parastylid-paraconid and entoconid-entostylid. The p4 is also long with an anteroposteriorly expanded metaconid and open lingual valleys. The tooth is pretty molariform. The lower molar has weak lingual ribs and anterior transverse flange. The median basal pillar is present.

Discussion

The specimens with pronounced neck, weak median ribs, divergent styles and open second lingual valley are represented by one species. The specimens are differentiated from the other bovids of Lava by the pronounced neck. The crown pronounced neck was present in Selenoportax and Helicoportax (Pilgrim, 1937, 1939). The Selenoportax was a large sized Siwalik boselaphine of the upper Miocene (Khan et al., 2007; Khan, 2008). The Lava specimens is smaller than the Selenoportax and correspond very well with it dental morphology and size to Helicoportax. The preserved specimens were not well enough to allow identifying of a species exactly. Nevertheless, the metrical values of the specimens are close to H. tragelaphoides and Helicoportax cf. tragelaphoides is assigned for the specimens (Table V, Fig. 5).

Table V.- Comparative dental measurements (mm) of Helicoportax.Comparative material is taken from Pilgrim (1937, 1939). * studied specimens.

Taxon###Specimen###Length###Width###

H. sp###*PUPC 07/114 - M1###17###17###

H. cf.tragelaphoides###GSI 822-M1###15###15.5###

H. tragelaphoides###AMNH 29909 - M1###17###17###

H. tragelaphoides###AMNH 29945 - M1###15###15###

H. praecox###AMNH 19476 - M1###12.5###15.5###

H. praecox###AMNH 19998 - M1###13###13###

H. praecox###AMNH 29867 - M1###13###13###

H. sp###*PUPC 09/55 - p3###15###7###

H. sp###*PUPC 09/55 - p4###13###8###

H. sp###*PUPC 09/55 - m1###14###10###

H. praecox###AMNH 29858 - p3###9.5###6.5###

H. praecox###AMNH 19995 - m1###13.5###8.5###

H. cf.tragelaphoides###GSI 821-m1###13###9###

H. cf.tragelaphoides###GSI 821-p3###13###11###

Kubanotragus sp.

Referred material

PUPC 07/113 - a fragment of horn core.

Description

The horn core is broken at the base as well as at the apex. Overall, the size of the horn core is not much large (Fig. 6D). The horn core fragment has irregular longitudinal striae. Its cross section is circular and it is slightly twisted. The horn core fragment shows a faint curvature. The light torsion is present in the horn core fragment.

Discussion

The horn core is distinguished from other medium size bovids e.g. Gazella, Sivaceros, Helicoportax, Eotragus and Elachistoceras by having very deep longitudinal groove (Thomas, 1984). It is differentiated from Caprotragoides potwaricus by having less mediolateral compression, gently curved and not stout (Bibi and Gulec, 2008). The horn core morphology and size are pretty fit within the range of Kubanotragus, identified by Thomas (1984) from the Chinji Formation.

Subfamily Giraffinae Gray, 1821

Tribe Palaeotragini Pilgrim, 1911

Genus GIRAFFOKERYX Pilgrim, 1910

Giraffokeryx punjabiensis Pilgrim, 1910

Referred material

PUPC 07/115 - rp3; PUPC 07/116 - rp4; PUPC 07/117 - rM1; PUPC 09/59 - lm1; PUPC 07/118 - lm3; PUPC 09/56 - lm3; PUPC 09/57 - lm3.

Description

The material consists of isolated teeth. The p3 has the paraconid well separated from the parastylid (Fig. 6E). The premolar is in early wear, brachydont, rugose and molarized. The metaconid extend forwards meeting the base of the paraconid. The crest joins to the metaconid and incorporates the entoconid. The entoconid is independent from the metaconid in early wear. A well developed furrow separates the hypoconid from the strong protoconid on the buccal side. The p4 is highly molariform (Fig. 6F). The metaconid is expanded lingually. The parastylid is thinner than in p3. The anterior lobe is large and closed lingually. The protoconid is connected to the hypoconid and tends to fuse with the entoconid lingually. The trigonid is distinguished by a furrow. The first molar is partially broken, simple with finely rippled enamel. The ectostylid and anterior fold are very weak but a well developed metastylid is present.

The upper molar has well developed styles and ribs but it has a weakly develop entostyle. The prtocone is angular and slightly constricted lingually (Fig. 6G). A cingulum is weakly developed. The hypoconulid of the third molar is elliptical forming a complete loop (Fig. 6H).

Discussion

The large sized brachydont teeth with sculpture enamel make their inclusion to giraffids (Colbert, 1935). The Siwalik giraffids are divided into two groups, one comprises the large forms of the upper Miocene and the other one comprises the small forms of the middle Miocene (Akhtar and Sarwar, 1987). The small sized Siwalik forms are

Giraffokeryx and Giraffa (Bhatti, 2005). The teeth display the character of Girafokeryx found in the Middle Miocene of the Siwaliks (Colbert, 1935; Geraads and Aslan, 2003; Geraads et al., 1995). The transverse orientation of the crests is observed in the specimens which is only found in the Siwalik G. punjabiensis and the specimens are assigned to G. punjabiensis. The species is more primitive than the Eurasian middle Miocene giraffids, having all crests of transverse orientation (Geraads and Aslan, 2003). Dimensions of the teeth are very similar to those of the Siwalik middle Miocene giraffids (Colbert, 1935; Bhatti et al., 2007). However, the variation in size may have some biostratigraphic value (Table VI, Fig. 7).

Table VI.- Comparative dental measurements (mm) of G. punjabiensis. * studied specimens.

Taxon###Specimen###Length###Width

. punjabiensis *###PUPC 07/115 - p3###25###14

G. punjabiensis *###PUPC 07/116 - p4###23.6###14

G. punjabiensis *###PUPC 07/117 - M1###28###28

G. punjabiensis *###PUPC 07/118 - m3###35.5###15

G. punjabiensis *###PUPC 09/56 - m3###26###cf. 13

G. punjabiensis *###PUPC 09/57 - m3###36###17

G. cf. punjabiensis###m3###34###17

(Geraads et al., 1995)###

G. cf. punjabiensis###p4###22.5###15.3

(Geraads et al., 1995)###

G. cf. punjabiensis###p4###21###13

(Geraads et al., 1995)###

G. punjabiensis###PUPC 02/12 - m3###34###18

(Bhatti, 2005)###

G. punjabiensis###PUPC 02/19 - m3###30###19

(Bhatti, 2005)###

G. punjabiensis###PUPC 02/06 - p4###23###14

(Bhatti, 2005)###

G. punjabiensis###AMNH 19317 - m3###37###18

(Colbert, 1935)###

G. punjabiensis###AMNH 19849 - m3###35###15

(Colbert, 1935)###

G. punjabiensis###AMNH 19587 - p4###24###15

(Colbert, 1935)###

G. punjabiensis###AMNH 19475 - M1###22###24

(Colbert, 1935)###

G. punjabiensis###AMNH 19587 - p3###20.5###12

(Colbert, 1935)###

G. punjabiensis###AMNH 19587 - m3###37###17

(Colbert, 1935)###

G. punjabiensis###AMNH 19472 - M1###23###22

(Colbert, 1935)###

Order Proboscidea Illiger, 1811

Family Deinotheriidae Bonaparte, 1845 Genus DEINOTHERIUM Kaup, 1829

Deinotherium cf. pentapotamiae

Referred material

PC-GCUF 09/29 - m1 (length, 26; width I ridge

plate, 17; width II ridge plate 20).

Description

The tooth is complete, isolated, unworn, brachydont and excellently preserved. The tooth is bilophodont with the contact facets (Fig. 6I). The crown of the tooth is elongated antero-posteriorly. The tubercles are present anterior to the first ridge plate. The posterior ridge plate is larger transversely than the anterior one. The ridge plates are concave anteriorly and convex posteriorly. The transverse valley between the first and the second ridge plate does not have any cingular tubercle. The cingulum surrounds the tooth anteriorly, buccally and posteriorly but it is prominent posteriorly where it forms a talon ridge enclosing a shallow valley.

Discussion

The tooth is bilophodont, typically the family Deinotheriidae. The true molar in Deinotherium, is always bilophodont (Sarwar, 1977). Three species of Deintherium D. pentapotamiae, D. orlovii and D. indicum are recognized in the Siwaliks (Sarwar, 1977) . D. orlovii and D. indicum are comparatively larger than D. pentapotamiae. The studied molar is small in size and can be assigned D. pentapotamiae. Nevertheless, differences in the dentition are present but are generally of minor nature, which renders specific identification. Therefore, the specific identification upon isolated teeth is a matter of some uncertainty.

BIOSTRATIGRAPHY AND CONCLUSIONS

Eight mammalian taxa have been recognized from the locality exposed in the south-eastern of village Lava, Jhelum, northern Pakistan. The fauna mainly consists of ruminants and rhinos, but the other groups are rare. The faunal list may be compared with the other middle Miocene Siwalik localities.

Brachyopotherium fatehjangense is a common species of the Siwalik middle Miocene (Heissig, 1972; Chavasseau et al., 2006). Gaindatherium browni was defined in the Chinji Formation and its stratigraphic range is late middle Miocene to early upper Miocene (Heissig, 1972; Colbert, 1935). The dates of occurrence for the deinotherium taxa suggest the Kamlial and the Chinji formations (ca. 18 - 14 Ma) in the Siwaliks (Tassy, 1983; Sarwar, 1977). Deinotherium pentapotamiae was found in the middle Miocene of the Lower Siwaliks (Dhem, 1963; Sarwar, 1977). In the Chinji strata Listriodon pentapotamiae is the most common suid (Pickford, 1988). Several Lower Siwalik localities of the Chinji Formation exhibit the presence of Listriodon pentapotamiae and the species is widely distributed in the Middle Miocene (Pickford, 1988; Pickford and Morales, 2003).

Highly lophodonts Listriodon pentapotamiae is abundant in the Chinji succession and deposits of the similar age elsewhere in the Potwar Plateau (Pickford and Morales, 200).

The Palaeotraginae is documented sporadically in the Chinji Formation of the Lower Siwaliks (Colbert, 1934; Bhatti et al., 2007).

Giraffokeryx punjabiensis has already been mentioned several localities of the late Middle Miocene age (Bhatti, 2005), occupying a wide territory from Western Europe to India (Bohlin, 1926; Bosscha-Erdbrink, 1977; Gentry et al., 1999; NOW database 2003). The Deinotherium pentapotamiae and Listriodon pentapotamiae are characteristic markers of the Chinji Formation (14 - 11.2 Ma). Helicoportax is also represented of the upper Chinji Formation (Pilgrim, 1937, 1939).

The long lasting and wide spread Listriodon pentapotamiae in the Lava locality together with Giraffokeryx punjabiensis, Gaindatherium browni, Brachypotherium fatehjangense, Deinotherium pentapotamiae, and Dorcatherium sp. indicates a middle Miocene age of the Lava locality. The fauna is in favor of a late Middle Miocene age because the comparison of the material with several representatives of the fauna indicates a middle Miocene age (Pilgrim, 1939, 1937; Thomas, 1984; Heissig, 1972; Pickford, 1988). The dental characters of the Lava material indicate closer affinities to the middle Miocene forms of the species, suggesting a similar age for the locality. In combination with the knowledge of the

Dorcatherium, Gaindatherium browni and Brachypotherium fatehjangense temporal distribution in the Siwaliks, it is likely that the site is middle Miocene in age. The taxa of the site indicate a time range that overlaps with the species in the Chinji Formation correspond to MN6 and MN7/8 (Heissig, 1972; Colbert, 1935; Farooq, 2006; Pickford and Morales, 2003). A similar vertebrate community is present in the Chinji succession of the Lower Siwaliks (Heissig, 1972; Pickford, 1988; Pilgrim, 1937, 1939; Colbert, 1935; Sarwar, 1977).

Ecological implications are less ambiguous. Lithofacies suggest a fluvial depositional environment. Brachypotherium has often been considered a marsh and lake dweller (Geraads and Sarac, 2003). Deinotherium indicates a forested habitat (Kay and Heissig, 2001). Listriodon seems to be a browser by judging from its dentition (Pickford, 1988). The selenodonty of the ruminants (Helicoportax, Giraffokeryx, Dorcatherium) in the locality may be interpreted for fibrous food which may have been the swamp vegetation due to the depositional environment (Janis et al., 2002). Dorcatherium is rather indicative of swampy-paludal habitats (Kohler, 1993; Rossner and Mors, 2001). This association points a wooded lake margin, surrounded by a more steppe or savannah like landscape.

ACKNOWLEDGEMENTS

We are grateful to Mr. Imran (Lecturer of Chemistry in Punjab University) and Malik Sahib (Landlord of the Lava village) for their hospitality during our stay in the Lava village. We would also like to thank Adeeb (M. Sc. student, GC University, Faisalabad) for working on the photographs and Razzaq for his efficient help extended during the fieldwork.

REFERENCES

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Zoology Department, Government College University, Faisalabad 2 Palaeontology laboratory, Zoology Department, Punjab University, Quid-e-Azam Campus, Lahore 3Department of Meteorology, COMSATS Institute of Information Technology, Islamabad 4Zoology Department, Science College, Wahdat Road, Lahore Corresponding author: akbaar111@yahoo.ca 0030-9923/2011/0001-0061 $ 8.00/0 Copyright 2011 Zoological Society of Pakistan.
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Author:Khan, Muhammad Akbar; Akhtar, Muhammad; Khan, Abdul Majid; Iqbal, Mehboob; Samiullah, Khizar; Ghaffa
Publication:Pakistan Journal of Zoology
Article Type:Report
Geographic Code:9PAKI
Date:Mar 31, 2011
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