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Proamphibos (Bovini: Bovidae: Ruminantia) from Sardhok Pleistocene of Pakistan.

Byline: Muhammad Khaled Siddiq Muhammad Akbar Khan and Muhammad Akhtar

Abstract

Some new Proamphibos fossils have been reported from Quaternary basin near Sardhok village in Gujrat northern Pakistan. The Sardhok Proamphibos sample includes an opisthocranium maxillar and mandibular fragments and isolated dentition. The bovine is well known in the Pakistani Siwaliks during Pleistocene. Proamphibos was represented by a reasonable number of specimens suggesting that the niche probably was filled by the large bovines. The new specimens suggest wooded grassland habitat and faunal turnover after 2.5 Ma. The paper represents one of the best Proamphibos records ever found from the subcontinent Siwaliks.

Keywords: Bovidae Pleistocene of Pakistan Pakistani Siwaliks.

INTRODUCTION

The Siwalik deposits represent a rich Neogene fossil records in the world ranging in age from Early Miocene to Middle Pleistocene (Matthew 1929; Colbert 1935; Pilgrim 1910

1913 1937 1939; Pilbeam et al. 1977 1979; Opdyke et al. 1979; Azzaroli and Napoleone 1982; Johnson et al. 1982 1985; Cande and Kent 1995; Barry et al. 1980 1982 1985 2002; Flynn et al.

1995 Flynn 2003; Flynn and Morgan 2005; Khan et al. 2009a b 2010 2011 2012a b). The Upper Siwaliks (ca. 3.4 0.6 Ma) traditionally subdivided into three lithological and faunal stages: Tatrot Pinjor and Boulder Conglomerates and it is better exposed in the Pabbi Hills Pinjor Stage (2.58 0.6

Ma) of Gujrat in northern Pakistan (Sarwar 1977; Hussain et al. 1992; Dennell et al. 2006 2008; Dennell 2008; Ghaffar et al. 2012). The Pabbi Hills fluvial deposits of Sardhok comprise the Pinjor Formation of the Upper Siwaliks (Shah 1977; Keller et al. 1977; Ranga Rao et al. 1988).

The Pinjor Formation of the type area in the Chandigarh region has been dated from 2.48 0.63 Ma by magnetostratigraphy (Kumaravel et al. 2005;

Nanda 2008). The Pinjor mammalian fauna ranging in age from 2.48 to 0.63 Ma is the youngest fauna of the Siwalik Group. The principal feature of

this community is continuing into the modern South Asian wildlife assemblages (Arif and Raza 1991; Arif and Shah 1991; Nanda 2002) characterized by dominance of herbivore community of woodland habitat with a few adapted for riverine galley forests.

The studied outcrops of the Sardhok village (Lat. 32 49' 39'' N: Long. 73 43' 51'' E) Gujrat district the Punjab province Pakistan is situated at

25 km SW of Jhelum city in the east of the Jhelum River and south of the Lahore Islamabad GT (Grand Trunk) road (Fig. 1). It is situated in the low altitude Pabbi Hills of the Upper Siwaliks yielding a very important mammalian fauna of Late Pliocene

Early Pleistocene age. The study area is characterized by brown to grayish-brown fine medium to coarse-grained sandstones with pebbles and large-scale cross stratifications brown mudstones pedogenic horizons and well-imbricated stratified conglomerates.

The Sardhok outcrops have not been explored by pioneer researchers (e.g. Falconer and Cautley 1849; Lydekker 1876 1880 1884; Pilgrim 1910 1913 1937 1939; Matthew 1929; Colbert 1935). A few specimens collected by Dr. Sarwar in the early 70ies and 80ies are mostly remained unpublished. The locality represents proboscideans perissodactyls and artiodactyls (Sarwar 1977; Dennell et al. 2006; unpublished data). At present the area in the vicinity of Sardhok has been thoroughly surveyed (Fig. 1). The specimens are comparatively scarce in these sites (numbered as SI SIX SXII SXIII S = Sardhok).

The new Proamphibos material originates from seven fossil sites: SII SIII SIV SV SVI SVIII and SX (Fig. 1). The systematic description of the new Proamphibos material is the focus of the article. The remains have been used to provide an indication of the probable age and paleoenvironment of these deposits.

MATERIALS AND METHODS

The present paper is a compilation of the Proamphibos fossil data newly recovered from the Sardhok Pleistocene of the Upper Siwaliks. The material is discovered by frequently visiting to macrofossil-yielding sites and part of the PhD thesis of the first author. The location of the excavated area in the Sardhok village is shown in figure 1. During the surveys a large area is covered over which fossils were present but at low densities and probably derived from more than one sedimentary unit. The collecting method is general. Some specimens were collected in gullies and a few in find-spots. The specimens excavated from these localities were generally in excellent condition with very little surface damage often complete and sometimes in articulation or anatomically adjacent to other specimens. The specimens found on the erosional surfaces were also well preserved particularly those that had not been exposed for long as on steep actively eroding slopes. The level surface specimens (fully exposed to sun and rain) are relatively poorly preserved. Almost all fossil specimens were found weathering out from or in situ within fine-grained yellow-buff sands and silts.

The spatial distribution of the fossil material was non-random and distributed in pockets a typical pattern of the Siwaliks (Raza et al. 2002; Barry et al. 2002). Many specimens were undiagnostic and few could be identified to cranial element and taxon at the generic or specific level. Measurements are expressed in millimeters. Morpho-metric features of the specimens are described and their systematic determination is discussed. The catalogue number on the specimen represents the collection year (numerator) and serial number (denominator) of that year (e.g. PUPC

11/19. Uppercase letters are used for upper dentition (e.g. M) and lower case for lower dentition (e.g. m). The crown terminology and measurement manners follow Pilgrim (1937 1939) Gentry et al. (1999) and BArmann and RAlssner (2011). The skull measurement manners follow Lei Zhu (2012).

SYSTEMATIC PALAEONTOLOGY

Mammalia Linnaeus 1758 Cetartiodactyla Montgelard Catzeflis and Douzery 1997

Pecora Linnaeus 1758 sensu Webb and Taylor 1980

Family Bovidae Gray 1821

Tribe Bubalina Pilgrim 1939

Genus PROAMPHIBOS Pilgrim 1939

Type species

Proamphibos lachrymans Pilgrim 1939.

Geographic distribution

Proamphibos is recorded from the Upper

Siwaliks of Pakistan India and Nepal (Pilgrim 1939; Khan et al. 2009a; Khan and Akhtar 2011).

Proamphibos kashmiricus Pilgrim 1939

Type specimen

A skull Geol. Surv. Ind. No. B 556.

Type locality

South east of Parmandal Samba tehsil

Jammu State India.

Diagnosis

Original diagnosis (Pilgrim 1939: 278).

Stratigraphic range

Upper Siwaliks.

New material

Opisthocranium: PUPC 10/87 female partial skull; Upper dentition: PUPC 66/80 rP4; PUPC 66/141 lP4; PUPC 68/54 rM1; PUPC 66/22 rM2; PUPC 67/295 rM2; Lower dentition: PUPC 10/4 rp3; PUPC 68/55 rp4; PUPC 69/475 lm1; PUPC 67/268 rm2; PUPC 67/116 a right mandible fragment with m2-3; PUPC 68/50 left mandible fragment with m2-3; PUPC 66/72 rm3; PUPC 67/283 rm3; PUPC 67/412 rm3; PUPC 68/196 rm3; PUPC 67/36 partial rm3; PUPC 67/302 partial rm3; PUPC 67/406 partial rm3; PUPC 68/59 partial rm3; PUPC 68/204 partial rm3; PUPC 66/144 lm3.

Locality

Sardhok Gujrat district Punjab Pakistan.

Description Opisthocranium

The skull is lacking the occipital condyles auditory bullae basioccipital area temporal fossa and most of the brain case (Fig. 2). The skull is not compressed dorso-ventrally. The frontal maxillary and premaxillary portion is preserved. The nasals are present. The frontals are perfect and narrow comparatively. It is slightly arched. The skull is hornless and therefore it is confirmed a female. The dentition is present but it is not clear for the morphological study (Fig. 2). The molars are lost their crown completely. The orbits are trapezoidal- shaped and partial damaged along the edges. The sutures separating the various bones are clearly visible; the interfrontal suture is also visible. The frontoparietal suture is partially preserved. The skull is slenderical dorsally and somewhat elongated. The brain case is partially preserved. The bregma separating the frontal from the parietal behind the horn cores is well represented.

The lambda between the parietal and the supraoccipital is missing. The frontals form two shallow depressions along the medial suture. The temporal ridges are absent. The orbit is laterally projected large and semi round (Fig. 2).

Upper dentition

The fourth premolars have half moon like outline (Fig. 3A B). The parastyle is moderately developed and the metastyle is strongly developed; between the two styles there is a prominent rib all along the crown height. The cementation is present labially. The cingulum is absent. The premolars are rugose and have three roots. The metastyle is slender rugose and covered by cement. The fossette with spur is crescent shape. The upper molars have four roots (Fig. 3C-E). The labial border of the anterior lobe is V-shaped and the posterior lobe is U-shaped. The molars have strongly developed parastyle mesostyle and metastyle. The paraconus and metaconus ribs are very strong. The vertical grooves are present between the mesostyle and the metaconus rib and between the paraconus rib and the mesostyle labially. The praeprotocrista is longer than postparacrista. The praehypocrista is smaller than the posthypocrista. The prae- and postmetacristae are of almost equal size.

The prae- and postfossette are wide and deep.

Lower dentition

The p3 is compressed transversely with a deep and wide anterolingual valley all along the crown height. There is a deep narrow valley posterolingually. The paraconid and protoconid occupies two-thirds of the premolar length (Fig. 3F). The p4 has a triangular outline; it has a deep anterolingual valley present all along the crown height. The paraconid bracketed by the anterolingual valley has in its lingual side a small stylid. In the posterolingual side of the premolar there is a deep valley (Fig. 3G). The circular-shaped fossettid is present by the fusion of the metaconid

and the entoconid. A shallow vertical groove is present posterolabially. The cementation is present and the cingulum is absent.

The lower molars are finely preserved with rugose enamel (Figs. 3H I; 4A-G). The protoconid hypoconid paraconid metaconid and entoconid are excellently preserved. The shallow vertical groove separating the protoconid from the protostylid is present antero-labially. The metastylid is heavy and looks like slender. The fossettids with spurs are crescent shape. There is a basal expansion of the metastylid. The ectostylid is heavy and more close to the protoconid than the hypoconid. The prae- entocristid is smaller than the postentocristid. The deep vertical groove narrow at the base and broad towards the top is present between entostylid and entoconid. The m3 is a three-lobed tooth with a strongly developed parastylid. The talonid of m3 is present posteriorly (Fig. 4). The measurements of the teeth are given in Table I.

Comparison

The absence of temporal lines is an indication of the absence of horn cores. The studied skull differs from the respective ones of Bison Bubalus Bos Leptobos and male Proamphibos in the absence of strong temporal lines (Martinez-Navarro et al. 2007). The weak or absent temporal lines of the skull are the features found in Bucapra Hemibos and female Proamphibos. Bucapra differs from the studied skull in having a narrow and high skull. The small mastoid width of the skull associates it to Proamphibos in distinction to Hemibos (Pilgrim 1937 1939). Two species of Proamphibos P. lachrymans and P. kashmiricus are present in the Siwaliks. The skull surface is not rugose the rugosity is normally absent in female P. kashmiricus (Pilgrim 1939). Morphometrically the studied skull resembles with that of a female P. kashmiricus (Table I; Fig. 2). The absence of the temporal lines indicates that the specimen belongs to a hornless female of P. kashmiricus.

The features width of the skull at the mastoid process and the narrowness of the skull fit pretty well with the previously described female skull of P. kashmiricus (Table II with comparative measures).

The teeth differ from Alcelaphini in having anterior transverse flange and ectostylid which are absent in alcelaphines (Gentry 1978). The teeth differ from Boselaphini in having cement disappearance of enamel wrinkling and enlargement of entostyle. The teeth associate with Bovini in having large size anterior transverse flange hypsodonty strong entostyle/ectostylid strong

Table I.- Comparison of the cheek teeth (mm) of Proamphibos. the studied specimens. Referred data are taken from Pilgrim (1939) Akhtar (1992) and Khan et al. (2009a).

Taxa###Specimen inventory No.###Nature###Length###Width###W/L

P. kashmiricus###PUPC 66/80###rP4###21.25###21.10###0.98

###PUPC 66/141###lP4###19.85###21.70###1.09

###PUPC 68/54###rM1###30.35###19.10###0.62

###PUPC 66/22###rM2###35.60###26.05###0.73

###PUPC 67/295###rM2###30.25###19.25###0.63

###PUPC 68/55###rp4###23.7###9.250###0.39

###PUPC 69/475###lm1###30.75###14.10###0.45

###PUPC 67/116###rm2###27.70###13.30###0.48

###rm3###36.00###13.65###0.37

###PUPC 67/268###rm2###32.75###18.25###0.55

###PUPC 68/50###lm2###32.30###16.05###0.49

###lm3###42.10###16.05###0.38

###PUPC 66/72###rm3###37.80###14.85###0.39

###PUPC 67/36###rm3###35.30###14.90###0.42

###PUPC 67/283###rm3###44.35###17.25###0.38

###PUPC 67/302###rm3###35.80###16.15###0.45

###PUPC 67/406###rm3###37.70###16.80###0.44

###PUPC 67/412###rm3###44.50###16.25###0.36

###PUPC 68/59###rm3###41.50###13.70###0.33

###PUPC 68/196###rm3###42.20###15.90###0.37

###PUPC 68/204###rm3###35.05###13.20###0.37

###PUPC 66/144###lm3###31.75###15.05###0.47

###GSI B561###P4###19.00###25.00###1.31

###M1###26.00###26.00###1.00

###M2###31.00###27.00###0.87

###PUPC 84/27###P4###18###19.5###1.08

###M1###27###25###0.92

###M2###32###23###0.71

###PUPC 72/57###P4###20###23###1.15

###GSI B561###m3###31###24###0.77

P. sp.###PUPC 69/641###P4###21###24###1.14

###M1###24###26###1.08

###M2###30###27###0.90

P. lachrymans###PUPC 68/79###m1###25###16###0.64

###m2###28###16###0.57

###m3###42###16###0.38

###B810###m1###22.5###18###0.80

###m2###24.5###18###0.73

###m3###34.5###18###0.52

###B811###m1###21###16###0.76

P. dhokawanensis###PUPC 69/351###m1###29###14###0.48

###m2###35###15###0.38

###m3###39###15###0.38

Table II.- Comparison of the cranial measurements (mm) of female Proamphibos.

###P. kashmiricus###P. kashmiricus###Bucapra daviesii###P. kashmiricus

###female###female###(BMNH 36677###female

Description###(PUPC 71/57)###(GSI B817)###Rutimeyer 1878)###(PUPC 10/87)

###Khan and###Pilgrim (1939)###Pilgrim 1939###studied sample

###Akhtar (2011)

Breadth of skull at mastoid###200###164###160###138

Skull length###-###-###-###156.5

Cranial length###-###-###-###102

Width of the braincase###114###112###105###96

Basal length###-###-###-###333

Distance from fronto-nasal suture to fronto-###-###126###-###104

parietal suture

Premolare-prosthion###-###-###-###62

Short skull length###-###-###-###263.6

Oral palatal length###-###-###-###174.4

Basion-palatinoorale###-###-###-###155.5

Lateral length of the premaxilla###-###-###-###72.4

Lateral facial length###-###-###-###135.8

Length of the braincase###-###-###-###112

Greatest inner length of the orbit###-###-###-###62

Greatest inner height of the orbit###-###-###-###62

Height of nuchal crest###-###-###-###48.7

Distance between orbital ridge and frontal suture###-###-###-###66

Length of frontal suture###-###-###-###117.8

Distance between supra-orbital pit and orbital###-###-###-###28.3

ridge

Distance between supra-orbital pits###-###82###-###68

Distance between supra-orbital pit and frontal###-###-###-###38

suture

Length of upper premolar series###-###62###63###48.6

Length of upper molar series###-###87###85###78.5

style/stylid and prominent ribs (Pilgrim 1939; Khan et al. 2009a). The excessive antero-posterior compression confirms the specimens of Bovini (Figs. 3 4). The excessive antero-posterior compression of Bovini molars has produced median ribs of extraordinary strength. The quadrate shape of the upper molars is a feature of the bovine kinds Proleptobos and Proamphibos and equally some elongation has taken place in Leptobos Hemibos and in a much less degree in Bubalus. The morphometery of the dental remains fit with

Proamphibos kashmiricus (Pilgrim 1939; Khan and Akhtar 2011) and the material is referred to Proamphibos kashmiricus.

DISCUSSION

The fossil occurrence of Proamphibos along with Equus Cervus Bubalus and Elephas were recorded from the Sardhok outcrops. The genus Equus appeared in south Asia ca. 2.5 Ma (Lindsay et al. 1980). Equus sivalensis Interval-Zone or the Pinjor Stage is still treated as a single unit that lasts without internal subdivisions from 2.60.6 Ma. Elephas planifrons Interval-Zone is little later ca. 2.9 Ma. Equus and cervids with antlers entered arrived in the Subcontinent around the Gauss Matuyama boundary ca. 2.58 Ma (Barry et al. 1995). The stratigraphic range of the assemblage suggests the age dated between 2.6 and 0.6 Ma.

The hypsodont teeth are actual an adaptation for more fibrous or abrasive plants in more open and arid habitat (Van Valen 1960; Fortelius 1985; Janis and Fortelius 1988; Solounias et al. 1994; Fortelius and Solounias 2000). Damuth and Fortelius (2001) noted that hypsodonty might be result of overall dry environment or a regularly occurring dry season with other factors (Fortelius 1985; Janis and Fortelius 1988). The large body size hypsodonty and robust dentition in bovines indicate dry-season grasslands during Early Pleistocene. Proamphibos implies grassy woodlands penetrating open forests formed under drier conditions and opened vegetation. The bovines would live here together with proboscideans cervids and antilopes (Dennell et al. 2006 2008; Khan et al. 2011).

The present study suggests a major turnover in land mammals which occurred after 2.5 Ma indicating a change towards a cooler drier and more variable climate. Proamphibos is the grazer and inhabitant more-open areas near water. Many other species that were bound to forest and wet habitats (e.g. Tragoportax Selenoportax Pachyportax Dorcatherium Dorcabune) disappeared during the time interval (Khan et al. 2009b 2012b). The associated change has been observed in the Northern Hemisphere and Africa at about this time (2.5 Ma) (Montgelard et al. 1997; Behrensmeyer et al. 1997; Williams et al. 1999).

The faunal change has been observed in the outcrops nearby the Sardhok village in the Pleistocene. The macro-fauna is indicatives of climatic change in the Pleistocene leading to increase aridity cooling and variable seasonality. The large mammals might have adapted wooded grassland semiarid sandy plains and seasonal pool pond bank. The significant findings of the large hypsodont mammals may be related to the intensification of wooded grassland (Fortelius et al. 2002).

CONCLUSIONS

The large mammals have been noticed in the outcrops of Sardhok. The Sardhok outcrops were mainly characterized by the presence Proamphibos kashmiricus with other mammalian taxa like Equus Elephas Hemibos Hexaprotodon and Rhinoceros during the Pleistocene. These taxa indicate the open wooded grasslands ecosystem in the Pabbi Hills of Sardhok northern Pakistan. Proamphibos indicates open country forms (Graze or Browse) and support aridity in the Pleistocene. The habitat spectrum indicates grazing forms dominated over browsers and browsing-grazing forms.

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Author:Siddiq, Muhammad Khaled; Khan, Muhammad Akbar; Akhtar Abstract, Muhammad
Publication:Pakistan Journal of Zoology
Article Type:Report
Geographic Code:9PAKI
Date:Aug 31, 2014
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