ISSN 0031�0301, Paleontological Journal, 2014, Vol. 48, No. 1, pp. 79–89. © Pleiades Publishing, Ltd., 2014.Original Russian Text © M.A. Khan, M. Akhtar, 2014, published in Paleontologicheskii Zhurnal, 2014, No. 1, pp. 80–91.

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INTRODUCTION

The tribe Antilopini is known from the MiddleMiocene Siwaliks (Akhtar, 1992); in the terminalMiocene and basal Pliocene, the group became wide�spread in Europe, Asia, and Africa (Bouvrain, 1979;Gentry et al., 1999; Geraads and Gulec, 1999).Reduncine antelopes appear for the first time in thefossil record in the Kundrali Unit (Dhok Pathan,Middle Siwaliks, Pakistan), along with a species verysimilar to Prostrepsiceros vinayaki (Pilgrim, 1939;Thomas, 1984; Bibi et al., 2009). The earliest redun�cine records in Eurasia are known from the Siwaliks ofIndia and Pakistan and dated Late Miocene (Gentry,1992, 1997; Vrba, 1995; Bibi, 2011). There are threeSiwalik reduncine taxa, Kobus porrecticornis (previ�ously assigned to the genus Dorcadoxa), Sivacobuspalaeindicus, and Vishnucobus patulicornis, each ofwhich had close relatives in Africa (Pilgrim, 1939;Vrba and Haile�Selassie, 2006).

The genus Antilope Pallas, 1766, which is presentlywidespread in Asia, is recorded in the Pinjor andTatrot formations of the Upper Siwaliks (Pilgrim,1937, 1939; Akhtar, 1992), in the Omo Group ofAfrica (Gentry, 1985), and in the Late Pliocene ofGerakarou, Greece (Kostopoulos, 1998; Kostopoulosand Koufos, 2006). Antilope is the only known genuswith two species, A. subtorta from the Pliocene of theSiwaliks (Pilgrim, 1937, 1939; Akhtar, 1992) andA. cervicapra from the Pleistocene of the Siwaliks (Pil�grim, 1939; Chauhan, 2007).

The taxonomic position of Siwalik antelopesrelated to living Antilope remains uncertain. Actually,no effort has been made after Pilgrim (1939) to clarify

the taxonomic status of Siwalik antelopes. The newfossil material allows the study of morphological vari�ability of antelopes from the Upper Siwaliks. In addi�tion, based on this material, the presence of a newAntilope species is established, which along withalready known antelopes surviving in the arid hillyareas of the Indian subcontinent shows stable exist�ence of antelopes in the Late Neogene–Quaternary ofthis region.

Stratigraphy and Age

The village of Tatrot (32°59′ N, 73°21′ E) is situ�ated approximately 30 km west of the Jhelum city,Punjab Province, Pakistan (Fig. 1). In the vicinity ofthe village, which is located at the boundary betweenthe Dhok Pathan Formation and younger depositsknown as the Upper Siwaliks, landscapes are charac�terized by low hills and flat lowlands. The Upper Siwa�lik deposits of the Indian subcontinent range from theLate Pliocene to Middle Pleistocene and dated about3.4–0.6 Ma (Nanda, 2002; Dennell et al., 2006).Since Pilgrim’s (1910, 1913) studies, the Upper Siwa�liks is subdivided into three lithological and faunalstages: the Tatrot stage, Pinjor stage, and BoulderConglomerate. Lithologically, the three stages consistof a sandstone–mudstone series, followed by the Par�mandal Sandstone and Boulder Conglomerate For�mation, which is the uppermost lithostratigraphicunit. The Tatrot Formation is the oldest part of theUpper Siwalik depositional sequence, consisting ofpale pinkish, orange, brown clay, brown and gray silt�stones and shales, greenish gray, fine to medium�

Antelopes (Mammalia, Ruminantia, Bovidae) from the Upper Siwaliks of Tatrot, Pakistan,

with Description of a New SpeciesM. A. Khan and M. Akhtar

Paleontology Laboratory, Department of Zoology, University of Punjab, Lahore, Pakistane�mail: akbaar111@yahoo.ca

Received November 28, 2012

Abstract—New antelope material from the Tatrot type locality of the Tatrot Formation, Upper Siwaliks,northern Pakistan, allows us to recognize four antelopes, i.e., Antilope intermedia sp. nov. Khan and Akhtar,Antilope subtorta Pilgrim, 1937, Antilope cervicapra (Linnaeus, 1758), and Kobus porrecticornis (Lydekker,1878). Comparisons with A. cervicapra and A. subtorta suggest that A. intermedia sp. nov. is closely related tothe black buck A. cervicapra. The new material from Tatrot confirms the idea that antelopes in the UpperPliocene Tatrot Formation were rather diverse.

Keywords: Antilope, Bovidae, Kobus, reduncines, Pliocene, Siwaliks

DOI: 10.1134/S0031030114010055

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Fig. 1. Map of the Potwar Plateau (left) indicating the studied area and generalized stratigraphic section (right) of the main Siwa�lik formations (after Behrensmeyer and Barry, 2005, modified; boundary dates after Dennell et al., 2008 and Nanda, 2008).

Conglo�merate

Pathan

grained sandstones and interbedding dark gray con�glomerates (Shah, 1980). The Tatrot Formation at thetype locality is on average 300 m thick.

In the previous studies, the Tatrot Formation wasassigned to the upper part of the Gauss Chron (Barryet al., 1982), that is, between 2.9 and 2.5 Ma on theBerggren et al. (1985) time scale. Hussain et al. (1992)proposed that the Tatrot Formation might be older,dated between 3.4 and 3.2, i.e., the lower part of theGauss; and Barry et al. (2002) dated this formation as3.5–3.3 Ma. Recently, Dennell et al. (2008) andNanda (2008) dated the upper and lower boundaries ofthe Tatrot Formation as 2.6–2.4 Ma and 3.4 Ma,respectively. Thus, the Tatrot Formation roughly cor�responds to the uppermost Pliocene (Fig. 1).

MATERIAL AND METHODS

The fossil material was obtained in the past decadesfrom the Upper Siwalik beds outcropping near the vil�lage of Tatrot; it is housed in the Paleontology Labora�tory the of Zoology Department, University of Pun�jab, Lahore, Pakistan. It includes a right horn core,mandible fragments, and isolated upper and lowermolars. The horn core is measured at its base. Theanteroposterior diameter (DAP) corresponds to thelarge diameter of the horn core base and it may benonparallel to the sagittal plane. The mediolateraldiameter (DT) is perpendicular to the DAP. The com�

pression index of the horn core is calculated as DT ×100/DAP.

The tooth length (L) and width (W) are measuredat the occlusal level. Lowercase and uppercase lettersdesignate the lower and upper teeth, respectively. Thetooth cusp terminology follows the nomenclature ofGentry (1994) and horn core terminology follows Pil�grim (1937) and Kostopoulos (1998). The measure�ments are given in mm.

Comparative material includes specimens fromGerakarou, northern Greece (GER), the AmericanMuseum of Natural History (AMNH), Natural His�tory Museum in London (BMNH), Geological Sur�vey of India (GSI), Geological Survey of Pakistan(GSP), Paleontological Collection of AgriculturalUniversity of Faisalabad (PC�AGUF), and PunjabUniversity Paleontological Collections (PUPC).

SYSTEMATIC PALAEONTOLOGY

Order Artiodactyla Owen, 1848

Family Bovidae GRAY, 1821

Subfamily Antilopinae GRAY, 1821

Tr i b e Antilopini Gray, 1821

Genus Antilope Pallas, 1766Antilope intermedia Khan et Akhtar, sp. nov.

E t y m o l o g y. From intermediate charactersbetween A. cervicapra and A. subtorta.

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H o l o t y p e. PCUP, no. 83/845, right horn corewith frontal fragment, dorsal part of orbit, and com�plete supraorbital notch; northern Pakistan, PotwarPlateau, Jhelum District, outcrops near the of villageTatrot (Upper Siwaliks); Upper Pliocene, Tatrot For�mation, 3.4–2.6 Ma.

D e s c r i p t i o n (Fig. 2). The holotype is a com�plete right horn�core with slightly damaged apex. It ismoderately long, positioned just above the orbit, andprobably diverges from its counterpart. Two horns,probably, several missed from each other. The horncore is weakly twisted and very closely spiraled. Thehorn core is circular in cross section at the base, lacksa keel, and narrows very slightly posteriorly; it is con�vex in anterior profile and concave in posterior profile.The supraorbital foramen is large, sunken in a deepand wide depression. It passes through the frontal intothe orbital cavity.

M e a s u r e m e n t s i n m m. Horn core lengthalong the anterior surface, ca. 175; horn core lengthalong the posterior surface, ca. 150; anteroposteriorhorn core diameter (DAP), 36.5; transverse diameterof horn core (DT), 34; index of compression, 93.1%(compression is almost absent).

C o m p a r i s o n. Two Antilope species, A. cervica�pra and A. subtorta, are known from the Upper Siwa�liks (Pilgrim, 1937, 1939). A. cervicapra differs fromthe new species in the strongly spiraled horn cores. InA. subtorta, horn cores are relatively weakly spiral andless twisted compared to the new species; it also distin�guished by the deep furrow extending along the horncore surface (Fig. 3).

The new species significantly differs from A. kou�fosi; the latter has weakly torsioned and open spiralhorn cores, which curve strongly backwards in the

middle part and recurve fairly upwards in a distal partto form a complex helix (Kostopoulos, 1998). Thehorn core spiraling is more open in A. koufosi than inA. intermedia sp. nov. (Fig. 3).

A. intermedia is intermediate between A. cervicapraand A. subtorta; the torsion of its horn core is greaterthan in A. subtorta and weaker than in A. cervicapra;the cross section is circular, close spiraling (whichmeans not swinging far out from the longitudinalstraight line between the base and tip of the horn core),without keels or a deep furrow; the supraorbital foram�ina are moderately large. A. intermedia shows a closerspiraling than in A. koufosi.

R e m a r k s. The DAP of A. cervicapra (BMNH,no. 37268) at the base is 31 mm and the DT is 29 mm,giving it an index of compression at DT × 100/DAP =93.5 % (Pilgrim, 1939). The compression index ofA. subtorta is the same as in A. cervicapra (Pilgrim,1937). The compression index can be used to relateA. intermedia sp. nov. with A. cervicapra and A. subtorta.

The spiraling and weak twisting (which meanstwisting or torsion of the axis or along the axis) in thehorn core under study allows its ascription to Antilo�pini (Pilgrim, 1939; Kostopoulos and Koufos, 2006)and the horn core compression index allows its inclu�sion in Antilope.

The horn core differs essentially from that ofknown Siwalik and other Eurasian genera. Thedimensions and morphology, i.e., weak torsion with�out a deep furrow and the closed spiraling suggest toassign it to a new species of the genus Antilope, A. inter�media sp. nov.

The spiral horned antelopes from Eurasia Anti�lospira, Spirocerus, and Gazellospira are also distin�

(a) (b) (c)

(d)

Posterior side

Anterior side

217

mm

Fig. 2. Antilope intermedia sp. nov., holotype PUPC, no. 83/845, right horn core: (a) anterolateral, (b) posterior, (c) anteromedialviews, and (d) cross section; Pakistan, Tatrot locality; Upper Pliocene, Tatrot Formation.

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PUPC 69/512 PUPC 83/845 AMNH 19989 GER�273

Fig. 3. Horn cores of antelopes (from left to right): Antilope cervicapra, specimen PUPC, no. 69/512; A. intermedia sp. nov., holo�type PUPC, no. 83/845; A. subtorta, specimen AMNH, no. 19989; and A. koufosi, specimen GER, no. 237. Scale bar, 30 mm.

guished by their tightly twisted horn cores with moreor less developed keels (Kostopoulos, 1998).

M a t e r i a l. Holotype.

Antilope subtorta Pilgrim, 1937

H o l o t y p e. AMNH, no. 19989, horn core frag�ment; northern Pakistan, Tatrot locality; UpperPliocene, Tatrot Formation, Upper Siwaliks beds.

D e s c r i p t i o n (Figs. 4a–4c). Specimen PUPC,no. 83/838, a mandible fragment is moderately thickand deep; p4 is badly damaged; the paraconid andparastylid are not preserved. The fourth premolarshows an open valley between the metaconid and theentoconid. The entoconid is somewhat distinct fromthe entostylid only at the crown apex. The hypoconidprojects laterally, with a moderately deep valley infront of it. The molars are narrow�crowned, moder�ately worn. The valleys are narrow; a small spurprojects into the anterior valley. The entostylids arewell developed and connected to the main body of thecrown. The median ribs are poorly developed and thegoat fold is strong (Fig. 4). The metastylid and ento�

stylid are poorly developed and stronger than themesostylid. The third molar has a very well developedhypoconulid with subovate fossette. The hypoconulidis not offset labially in this specimen.

C o m p a r i s o n. The presence of strong goat fold,the flat lingual wall of the lower molars, presence of theweak ribs between the stylids, and an expandedm3 hypoconulid with fossette associate the specimenwith Antilopini. Unlike specimen under study, thehypoconulid of the lower third molar in A. koufosi issubrounded and the lower molars of this species showweak goat folds (Kostopoulos, 1998). PUPC,no. 83/838 differs from A. cervicapra in the hypo�conulid with a fossette which is not offset labially. Thegoat fold is rather strong in A. cervicapra. SpecimenPUPC, no. 83/838 (Fig. 4) is very similar in morpho�logical and metrical characteristics to the paratype ofA. subtorta (Pilgrim, 1937: 812), originally knownfrom the Upper Siwaliks.

M a t e r i a l. Specimen PUPC, no. 83/838, a rightmandible with m1–m3 and posterior part of p4 fromthe Tatrot Formation, Upper Siwaliks beds of north�ern Pakistan.

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Antilope cervicapra (Linnaeus, 1758)

H o l o t y p e has not been designated; Recent spe�cies.

D e s c r i p t i o n (Figs. 4d–4f). The molars arepoorly worn, with shiny enamel. The goat fold isstrong in the molars. A weak ectostylid is present. Thefossettes are not transversely compressed. The hypo�conulid of the third molar is offset labially and lacks afossette.

C o m p a r i s o n. The mandible fragment (PC�AGUF, no. 01) is 88 mm long and somewhat deeperthan in A. subtorta. The mandible is more bulky androbust than that of A. subtorta. The molar series is53.6 mm long versus 44 mm in A. koufosi (Kostopou�los, 1998). The hypoconulid pattern of the specimendescribed is much more similar to that of A. cervicapraand different from that of A. subtorta and A. koufosi(Pilgrim, 1937; Kostopoulos, 1998). Morphometri�cally, the specimen in question (Figs. 4d–4f) resem�bles A. cervicapra and is assigned to A. cervicapra.

M a t e r i a l. Specimen PC�AGUF, no. 01, a rightmandible fragment with m1–m3 from the Tatrotlocality of northern Pakistan; Tatrot Formation,Upper Siwaliks beds.

Genus Kobus A. Smith, 1840

Kobus porrecticornis (Lydekker, 1878)

H o l o t y p e. GSIB229, frontal skull part withhorn cores; northern Pakistan, Tatrot locality; UpperSiwaliks, Tatrot Formation.

D e s c r i p t i o n (Figs. 5, 6). The horn core (spec�imen PUPC, no. 69/353) is incompletely preservedand broken at the apex (Fig. 5). The horn core curvesbackwards and compressed slightly mediolaterally.The mediolateral compression continues towards theapex. The cross section is subovate; the core is twistedslightly clockwise. The medial surface is convex at thebase, while the lateral surface is flat. The horn coresurface has continuous longitudinal ribs and groovedstrongly posteriorly by longitudinal furrows. A faintposterolateral keel is present (Fig. 5d).

The enamel of the upper molars is smooth andlacks wrinkles. The molars are hypsodont. SpecimenPUPC, no. 82/14 has a strong entostyle. The centralfossettes are complex in structure (with spurs) in bothupper molars. The lingual lobes of the upper molarsare constricted. The styles and ribs are strong (Fig. 6).The goat fold is large in the lower molars. The ecto�stylid is prominent. The labial lobes of the lowermolars are constricted. The lingual ribs and stylids arestrong (Fig. 6).

M e a s u r e m e n t s i n m m. Anteroposteriordiameter of the horn core base DAP = 30.6; DT = 25;the ratio DT × 100/DAP = 81.6%.

C o m p a r i s o n. The right horn core’s slightclockwise twist excludes the sample from Gazella.Nevertheless, the torsion may be observed in the horntip of some species of Gazella, but this does not extendto the horn cores (Pilgrim, 1939). The right horn coreis distinguished from Gazella rodleri in having clock�wise twist, because G. rodleri has an anticlockwise

(a)

(b)

(c)

(d)

(e)

(f)

Fig. 4. Mandible fragments of antelopes from the Tatrot locality, Tatrot Formation, Upper Pliocene: (a–c) Antilope subtorta, specimenPUPC, no. 83/838, right mandible fragment with m1–m3 and posterior part of p4; (d–f) Antilope cervicapra, specimen PC�AGUF,no. 01, right mandible fragment with m1–m3; (a, d) lingual, (b, e) labial, and (c, f) occlusal views. Scale bar, 10 mm.

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twist (Pilgrim, 1939). The horn core described isclearly differentiated from that of Antilope, Helicotra�gus, and Litocranius in the absence of strong torsion(Pilgrim, 1939; Thomas, 1984; Kostopoulos, 1998)and is close to the early Siwalik reduncines Kobus inregard to the degree of torsion, tendency to posterolat�eral keel, backward tilt, divergence, and sculpture. Thespecimen has a fainter keel than that of the earlyreduncines K. subdolus Gentry, 1980 and Kobus sp. 2 atLangebaanweg, and the Redunca aff. darti of Leh�mann, a character that could be explained by the ear�lier stage of antelope.

The overall direction, compression, slight twist,and sculpture of the horn core are very similar to thatof Siwalik K. porrecticornis (Lydekker, 1878; Pilgrim,1939; Thomas, 1980). This specimen differs from thecommon African reduncine in the greater transversecompression, less flattened lateral surface, less pro�nounced posterolateral angle, and less developed lon�gitudinal grooves on the posterior surface (Gentry,1997). The teeth under study are similar in morphol�ogy to K. porrecticornis (Table 1) and can be referred tothis species. The reduncine teeth studied in the Siwa�liks have the same dimensions size and hypsodonty asAfrican forms (Table 1), but differ in the stronger basalpillars and less pointed labial lobes (Gentry, 1997).The entostyle is present on M1 of early reduncines(Gentry, 1997).

R e m a r k s. Pilgrim (1939) referred the holotypeof this species (GSIB229) to Dorcadoxa porrecticornis

of the group Antilopini instead of Reduncini. Follow�ing Lydekker (1878), Pilgrim accepted that this speciesis related to the African genus Aepyceros and proposedthat Dorcadoxa was ancestral to extant African Aepyc�eros and could have been an early member of Antilo�pini, which migrated to Pakistan in the Miocene.Gentry (1970) proposed that this specimen (GSI, no.B229) is related to Reduncini. Later, Thomas (1980)and Gentry (1980) assigned this species to the genusKobus.

Thomas (1980) noted that Kobus porrecticornisappears between sandstones 6 and 7 at the base of theKundrali Unit in the type zone of Dhok Pathan (about8 Ma). Kobus porrecticornis from the Siwalik beds dis�plays features of early Reduncini. It was consideredmore primitive than African reduncines (Gentry,1997). Horn cores of K. subdolus, an early member ofAfrican Kobus, give evidence of probable evolutionaryadvantage relative to K. porrecticornis, since they aremore fully reduncine in characters, such as a less com�pressed cross section, more flattened lateral surface,better developed posterolateral keel, and weaker back�ward curvature in the basal part (Gentry, 1997).

M a t e r i a l. Specimen PUPC, no. 69/353, righthorn core fragment; specimen PUPC, no. 82/14, iso�lated right M1; PUPC, no. 82/13, isolated right M2;specimen; PUPC, no. 83/837, left mandible fragmentwith m1–m2; PUPC, nos. 88/03 and 02/137, rightmandible fragment with m2–m3; PUPC, no. 69/67,

(a) (b) (c) (d)

(e)

Fig. 5. Kobus porrecticornis, specimen PUPC, no. 69/353, right horn core fragment: (a) anterior, (b) medial, (c) lateral, and(d) posterior views; (e) cross section; Pakistan, Tatrot locality; Upper Pliocene, Tatrot Formation. Scale bar, 30 mm.

Fig. 6. Kobus porrecticornis from the Tatrot locality; Tatrot Formation, Upper Pliocene: (1) specimen PUPC, no. 82/14, right M1;(2) specimen PUPC, no. 82/13, right M2; (3) specimen PUPC, no. 83/837, left mandible fragment with m1–m2; (4) specimenPUPC, no. 88/03, right mandible fragment with m2–m3; (5) specimen PUPC, no. 83/816, left m3; (6) specimen PUPC,no. 69/67, right m3; (a) occlusal, (b) lingual, and (c) labial views. Scale bar, 10 mm.

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1а 2а 5а 6а

1b 2b5b

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isolated right m3; PUPC, no. 83/816, isolated left m3;all from the type locality.

DISCUSSION

Among living Bovidae, only the Asiatic speciesA. cervicapra belongs to Antilope. Lydekker (1885) cat�alogued a horn�core (BMNH, no. 37264) from theNarbada alluvium at Piparea under the name A. cervi�capra. Later, Lydekker (1886) ascribed to the samespecies another horn core fragment (specimen GSI,no. F100) from the Jamna alluvial beds and an isolatedsecond upper molar (specimen GSI, no. F287) fromBilla Surgam caves of Karnul District, India.

Pilgrim (1937) established a new species, A. sub�torta, based on a horn core fragment (holotypeAMNH, no. 19989), a cast of which is stored in the

Geological Survey of Pakistan. Horn cores of A. sub�torta have a clear longitudinal furrow and are consid�erably less twisted than that of A. cervicapra. Pilgrim(1937) also assigned of a second (specimen AMNH,no. 19990) and third (specimen AMNH, no. 19985)lower molars to this species. These teeth essentiallydiffer from lower molars of A. cervicapra in the greaterbreadth, the lower hypsodonty, and in the presence ofwell�developed median basal pillar. Later, Pilgrim(1939) excluded the third lower molar (AMNH, no.19985) from A. subtorta and transferred it to a new spe�cies, A. (?) planicornis, which was described based on aright horn core (BMNH, no. 37264) included byLydekker (1885) in catalogue under the name A. cervi�capra. According to Pilgrim (1939), the torsion of thehorn core is the same as in A. cervicapra, but less pro�nounced. The most significant distinctions of this

Table 1. Comparative measurements (mm) of the lower cheek teeth of the Tatrot antelopes; specimens examined in thepresent study are marked by *; published data are taken from Pilgrim (1937), Akhtar (1992), and Gentry (1997)

Taxon Specimen no. Tooth Length L Width W Ratio W/L

A. subtorta PUPC 83/838* rm1 13.8 9.6 0.69

rm2 19 11 0.57

rm3 24 11 0.45

PUPC 83/837 lm1 15 10.5 0.7

lm2 19 10.5 0.55

AMNH 19990 m2 16 9.5 0.59

AMNH 19985 m3 24.5 11.5 0.46

A. cervicapra PC�AGUF 01* rm1 15 10 0.66

rm2 17 10.5 0.61

rm3 21 10 0.47

PUPC 69/512 lm1 12 8.7 0.72

lm2 15 9.0 0.60

lm3 22 9.7 0.44

A. koufosi GER 148 lm1 14 9.7 0.69

lm2 17 10 0.58

lm3 25 9.5 0.38

K. porrecticornis PUPC 82/14* lM1 16.6 16.3 0.98

PUPC 82/13* lM2 17 15 0.88

PUPC 83/837* lm1 16.5 10.8 0.65

lm2 19 11.5 0.60

PUPC 88/03* rm2 17.4 9.6 0.55

rm3 23 9.0 0.39

PUPC 02/137* rm2 21 10.6 0.50

rm3 23.5 – –

PUPC 69/67* rm3 25.7 13 0.50

PUPC 83/816* lm3 28 12.5 0.44

K. aff. porrecticornis WM969/92 rM2 17.9 14.9 0.83

WM975/92 lm2 19.0 7.7 0.40

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specimen from the living species are the strong lateralcompression (the cross section is oval, anteroposteri�orly elongated) and the presence of a basal pillar in thelower third molar.

Antilope intermedia sp. nov. is identified as the thirdspecies of Antilope from the Tatrot locality of theUpper Siwaliks, which is established based on the spi�raling and torsion of the horn cores. Horn cores ofA. intermedia show torsion; horn cores of A. subtortaare loosely twisted and have a deep furrow, whereashorn cores of A. cervicapra are tightly twisted. Thecharacteristic spiraling is relatively close in A. subtortaand more open in A. cervicapra and A. koufosi (Pil�grim, 1937; Kostopoulos, 1998).

Vegetation and diet adaptations. The carbon iso�tope record demonstrates that, after 8.1 Ma, signifi�cant amounts of C4 grasses began to appear and, about6.8 Ma, floodplain habitats included extensive C4grasslands (Barry et al., 2002). Plant communitiesdominated by C3 forms were essentially reduced after7 Ma and communities dominated by C4 plants, whichmark open or grassy woodlands, appeared as early as7.4 Ma (Cerling et al., 1997; Barry et al., 2002). Thedata on antelopes support the reconstruction of envi�ronments mostly of grassy woodlands formed underdrier conditions and open landscapes and are thusconsistent with current environmental interpretationsfor the Late Pliocene of Eurasia and Africa.

Blackbuck (A. cervicapra) usually lives in herds of20–30 to several hundred, preferring open plains cov�ered with scrub and sometimes penetrating by thin for�ests (Geist, 1974; Prater, 1971). Horn cores of A. inter�media sp. nov. are rather strongly twisted on their axiswith close spiraling, suggesting similar function withthat of blackbuck. Horn cores of A. intermedia are notmore robust than in the living Indian blackbuck, but inEuropean Miocene spiral�horned antelopes (e.g.,Palaeoreas lindermayeri), horn cores can be robust andthick, implying intense use in intraspecific combat(Roberts, 1996; Geraads et al., 2003). Horns of thiskind are not found in smaller�sized and less sociableinhabitants of dense woods and completely open land�scapes (Geist, 1974; Janis, 1986). Antelopes lived herealong with reduncines and bovines of the Siwaliks(Dennell et al., 2006; Khan, 2007; Khan et al., 2008),suggesting that Tatrot provided an extremely diversespatiotemporal habitat structure, with a wide variety ofniches with browsing and mixed feeding.

P a l e o b i o g e o g r a p h y a n d p a l e o c l i �m a t e. Kobus porrecticornis is a prominent element ofthe Tatrot fauna, represented by a large number ofspecimens. It is considered the earliest reduncine,recorded from the Upper Middle Siwaliks of Pakistan(Thomas, 1980; Gentry, 1997; Bobe and Eck, 2001;Vrba and Haile�Selassie, 2006). Kobus aff. porrecticor�nis has been reported from the Kenyan Mpesida Bedsand the Lukeino Formation in Kenya (Thomas, 1979;Gentry, 1980; Vrba, 1995). The most ancient redun�cine records in Eurasia, in particular, in the Siwaliks

are dated Late Miocene of Dhok Pathan and LatePliocene of Tatrot (Vrba, 1995; Vrba and Haile�Selassie, 2006).

As Bibi et al. (2009) and Bibi (2011) noted, LatePliocene links between southern Asia and Africa southof Sahara are apparent in the shared presence in bothterritories of bovid taxa, such as Kobus porrecticornis.At the same time, three endemic Antilope species,A. intermedia, A. cervicapra, and A. subtorta, arerestricted to southern Asia and there is no evidence ofthe three species outside this continent. The subfami�lies Bovinae and Antilopinae became more diverseduring the Late Pliocene in the Siwaliks, while thetribe Boselaphini became rare in the Late Pliocene ofthe Siwaliks. The rarity of boselaphines in the Tatrotlocality suggests that the niche was probably filled bybovines and antilopines.

The evolution of the large body size and robustdentition in bovines was driven by climatic and envi�ronmental changes, i.e., intensification of seasonalaridity in southern Asia during the early Late Miocene(Bibi, 2007). Grasslands expanded at the expense ofwoodland habitats after the Early Pleistocene, therebyreducing arboreal habitats and browsing opportunities(Dennell et al., 2006). Researchers propose a strongnegative relationship between hypsodonty and meanannual precipitation, so that the high�crowned teethare indicative of a decease in rainfall (Damuth et al.,2002; Fortelius et al., 2002, 2006; Bibi, 2007).

The abundance of antilopines in the Tatrot localityindicates a mixture of woodland and grassland biomes.The taxonomic composition of Tatrot bovines indi�cates woodland and grassland habitats during the LatePliocene in the Upper Siwaliks (Barry et al., 1982,1995; Hussain et al., 1992; Dennell et al., 2006). Therarity of boselaphines in the Tatrot locality clearlyreflects their ecological differences. A closed habitathas been inferred for Late Miocene boselaphines(Khan et al., 2009). These habitats might have notbeen available for boselaphines to flourish in Tatrot.Reduncines and Antilope inhabit savanna, woodland,and grassland. In the Pliocene, the climate becamedrier, promoting the development of grasslands inplace of the contracting woodland (Basu, 2004), sug�gesting that more open habitats prevailed in Tatrot.

CONCLUSIONS

The Tatrot outcrops of the Late Pliocene UpperSiwaliks are characterized by the presence of fourantelope species, along with several other mammaliantaxa, including Equus sivalensis, Elephas planifrons,Hemibos triqueticornis, Hexaprotodon sivalensis, andRhinoceros sivalensis. These taxa indicate the dry�sea�son grasslands ecosystem in Tatrot. The genus Antilopeindicates open country forms and supports dryingtrends in the Late Pliocene. The faunal elements likeantelopes became dominant in Tatrot at the expense ofmost of the browsing and browsing–grazing forms.

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ACKNOWLEDGMENTS

We are grateful to S.M. Raza (Northeastern OhioUniversity College of Medicine Rootstown, USA andHigher Education Commission of Pakistan) for helpand encouragement to complete the present work.M.A. Khan would like to express gratitude to G. Röss�ner, who helped arranging the general structure andwording of the manuscript and to D.S. Kostopoulosfor his useful comments and linguistic support on themanuscript. We are also thankful to D. Geraads for hisbriefly thoughtful critique on the manuscript and toMuhammad Ali and Ali Hussain for providing us withmaterial from the Indian blackbuck to use it for ourcomparative study.

REFERENCES

Akhtar, M., Taxonomy and Distribution of the SiwalikBovids, Unpubl. PhD Thesis, Lahore: Univ. Punjab, 1992,pp. 1–372.

Barry, J.C., Lindsay, E.H., and Jacobs, L.L., A biostrati�graphic zonation of the middle and upper Siwaliks of thePotwar Plateau of northern Pakistan, Palaeogeogr., Palaeo�climatol., Palaeoecol., 1982, vol. 37, pp. 95–130.

Barry, J.C., Morgan, M.E., Flynn, L.J., et al., Patterns offaunal turnover and diversity in the Neogene Siwaliks of north�ern Pakistan, Palaeogeogr., Palaeoclimatol., Palaeoecol., 1995,vol. 115, pp. 209–226.

Barry, J., Morgan, M., Flynn, L., et al., Faunal and envi�ronmental change in the Late Miocene Siwaliks of northernPakistan, Palaeobiology, 2002, vol. 28, pp. 1–72.

Basu, P.K., Siwalik mammals of the Jammu Sub�Himalaya,India: An appraisal of their diversity and habitats, Quatern.Int., 2004, vol. 117, pp. 105–118.

Behrensmeyer, A. and Barry, J., Biostratigraphic surveys inthe Siwaliks of Pakistan: A method for standardized surfacesampling of the vertebrate fossil record, Palaeontol. Elec�tron., 2005, vol. 8, no. 1, pp. 1–24.

Berggren, W.A., Kent, D.V., Flynn, L.J., and Van Couver�ing, J.A., Cenozoic geochronology, Geol. Soc. Am. Bull.,1985, vol. 96, pp. 1407–1418.

Bibi, F., Origin, paleoecology, and paleobiogeography ofearly Bovini, Palaeogeogr., Palaeoclimatol., Palaeoecol.,2007, vol. 248, pp. 60–72.

Bibi, F., Mio�Pliocene faunal exchanges and African bioge�ography: The record of fossil bovids, Plos One, 2011, vol. 6,no. 2, p. e16688.

Bibi, F., Bukhsianidze, M., Gentry, A.W., et al., The fossilrecord and evolution of bovidae: State of the field, Palaeon�tol. Electron., 2009, vol. 12, no. 3, pp. 1–11.

Bobe, R. and Eck, G.G., Responses of African bovids toPliocene climatic change, Palaeobiology, 2001, vol. 27,pp. 1–47.

Bouvrain, G., Un nouveau genre de Bovide de la fin duMiocene, Bull. Soc. Géol. Fr., 1979, vol. 21, no. 4, pp. 507–511.

Cerling, T.E., Harris, J.M., Macfadden, B.J., et al., Globalvegetation change through the Miocene/Pliocene bound�ary, Nature, 1997, vol. 389, pp. 153–158.

Chauhan, P.R., Large mammal fossil occurrences and asso�ciated archaeological evidence in Pleistocene contexts ofpeninsular India and Sri Lanka, Quatern. Int., 2007,vol. 192, pp. 20–42.Damuth, J.D., Fortelius, M., Andrews, P., et al., Recon�structing mean annual precipitation based on mammaliandental morphology and local species richness, J. Vertebr.Paleontol., 2002, vol. 22, suppl. no. 48A, pp. 1–56.Dennell, R., Coard, R., and Turner, A., The biostratigraphyand magnetic polarity zonation of the Pabbi Hills, northernPakistan: An Upper Siwalik (Pinjor Stage) Upper Pliocene�Lower Pleistocene fluvial sequence, Palaeogeogr., Palaeo�climatol., Palaeoecol., 2006, vol. 234, pp. 168–185.Dennell, R.W., Coard, R., and Turner, A., Predators andscavengers in early Pleistocene of southern Asia, Quatern.Int., 2008, vol. 192, pp. 78–88.Fortelius, M., Eronen, J., Jernvall, J., et al., Fossil mam�mals resolve regional patterns of Eurasian climate changeover 20 million years, Evol. Ecol. Res., 2002, vol. 4,pp. 1005–1016.Fortelius, M., Eronen, J., Liu, L.P., et al., Late Mioceneand Pliocene large land mammals and climatic changes inEurasia, Palaeogeogr., Palaeoclimatol., Palaeoecol., 2006,vol. 238, pp. 219–227.Geist, V., On the relationship of social evolution and ecol�ogy in ungulates, Am. Zool., 1974, vol. 14, pp. 205–280.Gentry, A.W., The Bovidae (Mammalia) of the Fort Ternanfossil fauna, in Fossil Vertebrates of Africa, Leakey, L.S.B.and Savage, R.J.G., Eds., London: Academic Press, 1970,vol. 2, pp. 243–323.Gentry, A.W., Fossil Bovidae (Mammalia) from Lange�baanweg, South Africa, Ann. S. Afr. Mus., 1980, vol. 79,pp. 213–337.Gentry, A.W., The Bovidae of the Omo Group deposits,Ethiopia, in Les faunes Plio�Pleistocene de la BasseVallee deOmo (Ethopia), vol. 1 Perissodactyles–Artiodactyles(Bovidae), Paris: CNRS, 1985, pp. 119–191.Gentry, A.W., The subfamilies and tribes of Bovidae, Mam�mal Rev., 1992, vol. 22, pp. 1–32.Gentry, A.W., The Miocene differentiation of Old WorldPecora (Mammalia), Histor. Biol., 1994, vol. 7, pp. 115–158.Gentry, A.W., Fossil ruminants (Mammalia) from theManonga Valley, Tanzania, in Neogene Paleontology of theManonga Valley, Tanzania, Harrison, T.J., Ed., New York:Plenum Press, 1997, pp. 107–135.Gentry, A.W., Roessner, G.E., and Heizmann, E.P., Subor�der Ruminantia, in The Miocene Land Mammals of Europe,Rössner, G.E. and Heissig, K., Eds., München: Dr.Friedrich Pfeil, 1999, pp. 225–258.Geraads, D. and Gulec, E., On some spiral�horned ante�lopes (Mammalia: Artiodactyla: Bovidae) from the LateMiocene of Turkey, with remarks on their distribution,Paläontol. Z., 1999, vol. 73, nos. 3–4, pp. 403–409.Geraads, D., Spassov, N., and Kovachev, D., Palaeoreaslindermayeri (Wagner, 1848) (Mammalia, Bovidae) fromthe Upper Miocene of Bulgaria, and a revision of the spe�cies, Geodiversitas, 2003, vol. 25, no. 2, pp. 405–415.Hussain, S.T., Van den Bergh, G.D., Steensma, K.J., et al.,Biostratigraphy of the Plio�Pleistocene continental sedi�ments (Upper Siwaliks) of the Mangla�Samwal anticline,

PALEONTOLOGICAL JOURNAL Vol. 48 No. 1 2014

ANTELOPES (MAMMALIA, RUMINANTIA, BOVIDAE) 89

Azad Kashmir, Pakistan, Proc. Kon. Nederl. Akad. Wetensch.,1992, vol. 95, pp. 65–80.Janis, C., Evolution of horns and related structures, Discovery,1986, vol. 19, pp. 8–17.Khan, M.A., Taxonomic Studies on Fossil Remains ofRuminants from Tertiary Hills of Hasnot, Pakistan,PhD Theses, Lahore: Univ. Punjab, 2007, pp. 1–371.Khan, M.A., Khan, A.M., Iqbal, M., et al., Reduncine fos�sils from the Upper Siwaliks of Tatrot, J. Anim. Plant Sci.,2008, vol. 18, no. 1, pp. 50–52.Khan, M.A., Iqbal, M., Ghaffar, A., and Akhtar, M., Pro�amphibos (Bovini, Bovidae, Mammalia) from the TatrotFormation in the Upper Siwaliks of pakiStan, J. Anim. PlantSci., 2009, vol. 19, no. 2, pp. 104–107.Kostopoulos, D.S., Parastrepsiceros koufosi n. sp. (Mam�malia: Bovidae); note on the possible presence of a Paras�trepsiceros descendant in the latest Pliocene of northernGreece, Acta Zool. Cracov., 1998, vol. 41, no. 1, pp. 101–109.Kostopoulos, D.S. and Koufos, G., Pheraios chryssomallos,gen. et sp. nov. (Mammalia, Bovidae, Tragelaphini), fromthe Late Miocene of Thessaly (Greece): Implications fortragelaphin biogeography, J. Vertebr. Paleontol., 2006,vol. 26, no. 2, pp. 436–445.Lydekker, R., Indian Tertiary and post�Tertiary Vertebrata:3. Crania of ruminants, Palaeontol. Ind., 1878, vol. 10,pp. 88–181.Lydekker, R., A revision of the antelopes of the Siwaliks,Geol. Mag., 1885, vol. 3, no. 2, pp. 169–171.Lydekker, R., Indian Tertiary and post�Tertiary Vertebrata:Siwalik Mammalia, Palaeontol. Ind. Ser., 1886, vol. 1(suppl.), no. 10, pp. 1–22.Nanda, A.C., Upper Siwalik mammalian fauna of India andassociated events, J. Asian Earth Sci., 2002, vol. 21, pp. 47–58.Nanda, A.C., Comments on the Pinjor mammalian faunaof the Siwalik Group in relation to the post�Siwalik faunasof Peninsular India and Indo�Gangetic Plain, Quatern. Int.,2008, vol. 192, pp. 6–13.Pilgrim, G.E., Notices of new mammalian genera and spe�cies from the Tertiaries of India, Calcutta, Rec. Geol. Surv.India, 1910, vol. 40, pp. 63–71.

Pilgrim, G.E., Correlation of the Siwaliks with mammalhorizons of Europe, Rec. Geol. Surv. India, 1913, vol. 43,pp. 264–326.

Pilgrim, G.E., Siwalik antelopes and oxen in the AmericanMuseum of Natural History, Bull. Am. Mus. Natur. Hist.,1937, vol. 72, pp. 729–874.

Pilgrim, G.E., The fossil Bovidae of India, Palaeontol. Ind.N. S., 1939, vol. 26, no. 1, pp. 1–356.

Prater, S., The Book of Indian Animals, Bombay: BombayNatur. Hist. Soc., 1971.

Roberts, S.C., The evolution of hornedness in female rumi�nants, Behaviour, 1996, vol. 133, pp. 399–442.

Shah, S.M., Stratigraphy and economic geology of CentralSalt Range, Rec. Geol. Surv. Pakistan, 1980, vol. 52, pp. 1–104.

Thomas, H., Les Bovides miocènes des rifts est�africains:Implications paléobiogéographiques, Bull. Géol. Soc. Fr.Sér. 7, 1979, vol. 21, no. 3, pp. 295–299.

Thomas, H., Les bovides du Miocène supérieur des couchesde Mpesida et de la formation de Lukeino (district de Bar�ingo, Kenya), in Proceedings of the 8th Pan�African Congressof Prehistory, Nairobi, 1977, Leakey, R.E.F. and Ogot, B.A.,Eds., Nairobi, 1980, pp. 82–91.

Thomas, H., Les bovides ante�hipparions des Siwaliksinférieurs (plateau du Potwar), Pakistan, Mém. Soc. Géol.Fr., 1984, vol. 145, pp. 1–68.

Vrba, E.S., The fossil record of African antelopes (Mamma�lia, Bovidae) in relation to human evolution and palaeocli�mate, in Paleoclimate and Evolution: With Emphasis onHuman Origins, Vrba, E.S., Denton, G.H., Partridge, T.C.,and Burkle, L.H., Eds., New Haven: Yale Univ. Press, 1995,pp. 385–424.

Vrba, E. and Haile�Selassie, Y., A new antelope, Zephyre�duncinus oundagaisus (Reduncini, Artiodactyla, Bovidae),from the Late Miocene of the Middle Awash, Afar Rift,Ethiopia, J. Vertebr. Paleontol., 2006, vol. 26, no. 1,pp. 213–218.

Translated by G. Rautian