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A NEW ISECTOLOPHID TAPIROMORPH (PERISSODACTYLA,MAMMALIA) FROM THE EARLY EOCENE OF PAKISTANAuthor(s): M. C. MAAS, S. T. HUSSAIN, J. J. M. LEINDERS, and J. G. M. THEWISSENSource: Journal of Paleontology, 75(2):407-417. 2001.Published By: The Paleontological SocietyDOI: http://dx.doi.org/10.1666/0022-3360(2001)075<0407:ANITPM>2.0.CO;2URL: http://www.bioone.org/doi/full/10.1666/0022-3360%282001%29075%3C0407%3AANITPM%3E2.0.CO%3B2

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407

J. Paleont., 75(2), 2001, pp. 407–417Copyright q 2001, The Paleontological Society0022-3360/01/0075-407$03.00

A NEW ISECTOLOPHID TAPIROMORPH (PERISSODACTYLA, MAMMALIA)FROM THE EARLY EOCENE OF PAKISTAN

M. C. MAAS,1,2 S. T. HUSSAIN,3 J. J. M. LEINDERS,4 AND J. G. M. THEWISSEN1

1Department of Anatomy, Northeastern Ohio Universities College of Medicine, P.O. Box 95, Rootstown 44272,2Department of Anthropology and Laboratory of Vertebrate Paleontology, University of Texas, Austin 78712,

3Department of Anatomy, Howard University, Washington D.C. 20059, and4Open University of the Netherlands, P.O. Box 2960 6401 DL Heerlen, The Netherlands

ABSTRACT—A new genus and species of tapiromorph, Karagalax mamikhelensis, is described from the Eocene Mami Khel Formationof northwest Pakistan. The new species is known from adult and juvenile dentitions, juvenile skulls, and partial postcrania. It is themost primitive perissodactyl yet reported from Indo-Pakistan. The morphology of its lophodont molars indicates that Karagalax is atapiromorph, and it is here included in the primitive family Isectolophidae. Karagalax is more derived (more lophodont) than NorthAmerican isectolophids Systemodon and Cardiolophus or the Asian early Eocene Orientolophus and Homogalax wutuensis, and moreprimitive (less lophodont) than North American Homogalax and Isectolophus. It is distinct from the poorly known and enigmatic Indianisectolophid Sastrilophus. Karagalax lacks any derived features of the Deperetellidae, Helaletidae or Lophialetidae, including Kalakotia,a primitive lophialetid from the middle Eocene of northwest India. The partial postcrania of Karagalax, which include fragmentaryhumeri, femora, ulnae, tibiae and metapodials, show a combination of primitive and derived features and suggest that it was morecursorial than other basal tapiromorphs for which postcrania are known.

A provisional analysis of the phylogenetic positions of Karagalax and Kalakotia supports the hypothesis that primitive perissodactylsdispersed to Indo-Pakistan, most probably by way of continental Asia. The evolutionary position of Karagalax is consistent with anearly Eocene age for H-GSP Locality 300, as argued previously on the basis of other mammals.

INTRODUCTION

TWENTY YEARS ago a small but intriguing assemblage of earlyTertiary mammals was discovered at Howard University-

Geological Survey of Pakistan (H-GSP) Locality 300, locatedsouth of the small settlement of Barbora Banda (sometimesknown as Barbara Banda) and west of Banda Daud Shah in KarakDistrict (formerly part of Kohat District) of the North West Fron-tier Province of Pakistan. H-GSP Locality 300 also is known asBarbora Banda I; a second fossiliferous area, designated BarboraBanda II, is located within half a kilometer of Barbora Banda Iand is considered part of the same deposit (Wells, 1983). Thefossils from both areas comprise the Barbora Banda fauna.

The sedimentology and fossil occurrences at H-GSP Locality300 are described in detail by Wells (1983) and Thewissen et al.(1983, 1997). The fossils occur in granulestone conglomeraticlenses in the redbed unit of the Mami Khel Formation (Meissneret al., 1974; Pivnik and Wells, 1996) (also referred to as lowerKuldana Formation) and apparently accumulated as the result ofa catastrophic flashflood (Wells, 1983). Previously described taxafrom the Barbora Banda fauna include three rodents (de Bruijnet al., 1982), an omomyid primate (Russell and Gingerich, 1987),and the dichobunid artiodactyl Diacodexis pakistanensis (Thew-issen et al., 1983; Russell et al., 1983; Thewissen and Hussain,1990). H-GSP Locality 300 was revisited in 1997 and 1998, andseveral additional mammals were discovered (Thewissen et al., inreview).

H-GSP Locality 300 is remarkable for the absence of endemicEocene Indo-Pakistani mammalian families and for the qualityand completeness of its most common fossil mammal, the prim-itive artiodactyl Diacodexis pakistanensis. The absence of endem-ic groups has led some to suggest that Barbora Banda representsa time previously unsampled in early Tertiary circum-Tethyanmammal history, and one critical in its biogeographic history(Thewissen et al., 1987). The new tapiromorph perissodactyl fromH-GSP Locality 300 described here is based on the most completematerial for the order yet reported from the early Tertiary of theIndian subcontinent. As such, it promises to expand significantlyour understanding of the diversity and evolutionary history ofbasal tapiromorphs.

Tapiromorphs were recently redefined by Froehlich (1996,

1999), based on an explicit phylogenetic analysis of basal peris-sodactyls. According to Froehlich, tapiromorphs are a stem-basedclade that includes tapirs plus rhinos (ceratomorphs) and thosetaxa more closely related to tapirs and rhinos than to horses andtheir relatives (hippomorphs). Froehlich’s study is basically inagreement with Hooker’s (1989) previous phylogenetic analysisof the basal perissodactyls, except for the conclusion that the earlyEocene European Cymbalophus (Hooker, 1984) and North Amer-ican Systemodon (5Hyracotherium tapirinum) are tapiromorphs.Hooker (1989, 1994) considered them to be basal equoids.

Most basal tapiromorphs are included in the Isectolophidae, aparaphyletic family (Froehlich, 1999, but see Hooker, 1989) witha Holarctic distribution. They are best known from North Americawhere they are represented by Cardiolophus (Gingerich, 1991),Homogalax, Isectolophus, and Systemodon (see Radinsky, 1963;Froehlich, 1996). Cymbalophus (Hooker, 1984), which is knownonly from isolated teeth and reconstructed jaw fragments, is theonly European representative of the family. Four isectolophid spe-cies are known from Asia. Orientolophus hengdongensis, whichincludes Homogalax namadicus (Dashzeveg, 1979, 1988; Ting,1995), is from the earliest Eocene of China and Mongolia (Ting,1993, 1995, 1998). Homogalax wutuensis and another, unnamed,isectolophid are from early Eocene deposits in the Wutu Basin ofChina (Chow and Li, 1965; Radinsky, 1965b; Ting, 1993, 1998;Tong and Wang, 1998). Ting (1993, 1998) reports that Homogalaxwutuensis is slightly less primitive than Orientolophus, which isconsidered more primitive than any other tapiromorph (Ting,1993; 1998; Froehlich, 1999). Sastrilophus dehmi, known onlyfrom a single damaged P3 and associated M1–3, is from the middleEocene of India (Sahni and Khare, 1971). Its phylogenetic posi-tion is debatable (Schoch, 1989; McKenna and Bell, 1997), butif it is an isectolophid, it is the only isectolophid other than Kar-agalax yet described from Indo-Pakistan.

Other than Sastrilophus, all tapiromorphs from the Eocene ofIndo-Pakistan are included in three more derived families: Helal-etidae, Deperetellidae, and Lophialetidae (Russell and Zhai,1987)—the late Eocene perissodactyl Indolophus from nearbyMyanmar is probably not a tapiromorph because its M1 metalophnearly terminates before it reaches the ectoloph (Schoch, 1989).Kalakotia simplicidentata (Ranga Rao, 1972), a species from the

408 JOURNAL OF PALEONTOLOGY, V. 75, NO. 2, 2001

FIGURE 1—Karagalax mamikhelensis n. gen. and sp. 1, H-GSP 98186,holotype in stereo occlusal view. Right and left maxillae with P3-M3.2, H-GSP 98186, holotype in left lateral view. Scale bars 5 1 cm.

middle Eocene Subathu Formation of India, is considered themost primitive of the endemic Indo-Pakistan forms (Hooker,1989). Although Kalakotia generally is classified as a lophialetid(Russell and Zhai, 1987; McKenna and Bell, 1997), Schoch(1989) argued that it is not a lophialetid because it lacks the di-agnostic U-shaped lophoid loop on P3–4 (see also Hooker, 1989).Aulaxolophus quadrangularis (Ranga Rao, 1972), Chasmother-ium mckennai (Sahni and Khare, 1972), and Schlosseria radinskyi(Sahni and Khare, 1972) are considered junior synonyms of Ka-lakotia simplicidentata (Russell and Zhai, 1987; Thewissen et al.,1987).

Dental terminology follows Hooker (1989, fig. 6.1). Institu-tional abbreviations: American Museum of Natural History(AMNH), Howard University-Geological Survey of Pakistan (H-GSP), Lucknow University Vertebrate Paleontology Laboratory(LUVP), Oil and Natural Gas Commission, Directorate of Geol-ogy, India (ONG), United States Geological Survey (USGS), andthe University of Michigan Museum of Paleontology (UM). H-GSP specimens are permanently deposited at the Geological Sur-vey of Pakistan, Stratigraphy/Paleontology Branch, Islamabad,Pakistan.

SYSTEMATIC PALEONTOLOGY

Order PERISSODACTYLA Owen, 1848Infraorder TAPIROMORPHA Haeckel, 1866Family ISECTOLOPHIDAE Peterson, 1919

KARAGALAX new genusType species.Karagalax mamikhelensis new species.Included species.Type species only.Differential diagnosis.Differs from Isectolophus and resem-

bles Homogalax, Cardiolophus, Systemodon and Orientolophusin retention of very small molar paraconules and metaconules andpresence of pre- and postcanine diastemata. Differs from Car-diolophus, Orientolophus, Systemodon and most Homogalax inmore molariform premolars (better development of P3 and P4 post-protocrista and P4 protolophid). Differs from Cardiolophus inhigher-crowned, less bunodont molars and premolars. Differsfrom Orientolophus in more lophodont molars (absence of me-tastylid or twinned metaconid, less notched hypolophid and pro-tolophid, cristid obliqua less lingually directed). Differs from Sas-trilophus in more molariform P3 and lingually deflected anteriormolar metacone. Differs from Lophialetidae except Kalakotia inabsence of complete, lophoid loop (pre- and postprotocristae) onP3–4. Differs from Kalakotia in retention of molar conules, lesslophodont molars (protolophid and hypolophid lower andnotched), and convex molar metacones.

Etymology.From ‘Karak,’ in reference to the Karak District,North West Frontier Province, Pakistan and Greek galax—milk,in reference to the early Eocene tapiromorph Homogalax and tothe nearly complete deciduous dentition of the referred specimens.

Karaglax mamikhelensis new speciesFigures 1–3; Table 1

Holotype.H-GSP 98186 right and left maxillae with P3-M3.Etymology.For Mami Khel Formation, North West Frontier

Province, Pakistan.Diagnosis.As for genus.Material examined.H-GSP 5139, left mandible with P1, P2

alveoli, P3-M3, and right mandible with P1 alveolus, P2-M3; H-GSP 5142, left Mx; H-GSP 5143, right Mx; H-GSP 5144, rightM3; H-GSP 5145, right M3; H-GSP 5223, juvenile skull with leftDP1 (roots only), DP2–4, M1, erupting M2, and right DP1 (roots),DP2–4, M1, erupting M2; H-GSP 5224, juvenile skull with right C1

(erupting), DP1–4, M1, erupting M2 and left C1, DP1–4, M1; H-GSP5225, left mandible with M1 (root), M2, posterior half M3; H-GSP5226, associated rostrum and mandible with right erupting I3, C1,

DP1, DP2–3 (damaged), left DP1–4, M1 (base of teeth only), eruptingM2, and right I3, C1, DP1(?), DP2–4 (damaged), M1, erupting M2,left C1, DP1?, DP3, DP4, trigonid M1, erupting M2; H-GSP 5229,right mandible with posterior root P3, damaged P4, M1; H-GSP5230, right M1 or 2; H-GSP 98157, left DP1; H-GSP 98159, distalfemur; H-GSP 98160, long bone fragment; H-GSP 98161, juve-nile right distal femur; H-GSP 98162, juvenile right femur; H-GSP 98163, distal humerus; H-GSP 98164, juvenile right proxi-mal tibia; H-GSP 98169, thoracic vertebra fragment; H-GSP98170, caudal vertebra ; H-GSP 98171, metapodial; H-GSP98172, left astragalus; H-GSP 98173, sesamoid; H-GSP 98174,proximal phalanx; H-GSP 98175, proximal phalanx; H-GSP98176, intermediate phalanx; H-GSP 98177, distal phalanx; H-GSP 98178, metapodial head; H-GSP 98179, juvenile left hu-merus; H-GSP 98180, juvenile right humerus; H-GSP 98181, dis-tal radius; H-GSP 98182, radius—epiphyseal plate; H-GSP98183, right proximal ulna; H-GSP 98184, distal tibia.

Occurrence.H-GSP Locality 300 (Barbora Banda I Locality),

409MAAS ET AL.—NEW TAPIROMORPH FROM PAKISTAN

FIGURE 2—Karagalax mamikhelensis n. gen. and sp. 1, H-GSP 5139, stereo occlusal view left P1, P2 alveoli, P3-M3 and right P1 alveolus, P2-M3; 2,H-GSP 5139, right lateral view; 3, H-GSP 5139, left lateral view. Scale bars 5 1 cm.

approximately 7 km west of Banda Daud Shah village and 1.5km south of the settlement of Barbora Banda (Latitude338159120N, Longitude 71889200E), Karak District, North WestFrontier Province, Pakistan; Mami Khel Formation, early Eocene.

Description.Three specimens preserve portions of the skull,including partial basicrania (H-GSP 5223 and H-GSP 5224), na-sals, and maxillae (H-GSP 5226). All three represent juvenile in-dividuals, and it is possible that the morphology of the skull hadnot reached its final proportions in these. Several bones are miss-ing from the skulls of these individuals, probably because they

were not fused in life to the skulls. The morphology of severalspecimens also was distorted post-mortem.

Based on a juvenile specimen (H-GSP 5226), the nasal incisureis deep, reaching posteriorly to the area over the canine (Fig. 3.3).The nasals roof this incisure and bridge the entire precanine di-astema, which is 12 mm in length. The anterior extent of thenasals cannot be determined because their tip is broken. Theerupting canine is separated from DP1 by a postcanine diastemaof 4.5 mm (H-GSP 5224).

The infraorbital foramen is located over the anterior root of P4,

410 JOURNAL OF PALEONTOLOGY, V. 75, NO. 2, 2001

FIGURE 3—Karagalax mamikhelensis n. gen. and sp. 1, H-GSP 5223,juvenile skull, ventral view, with left DP1 (roots only), DP2–4, M1, erupt-ing M2, and right DP1 (roots), DP2–4, M1, erupting M2; 2, H-GSP 5224,juvenile skull, dorsal view; 3, H-GSP 5226, associated rostrum andmandible, right buccal view showing right erupting I3, C1, DP1, DP2–3

(damaged), and right I3, C1, DP1(?), DP2–4 (damaged), M1, erupting M2;4, H-GSP 98172, left astragalus. Scale bars 5 1 cm.

and the anterior orbit in the adult is over the posterior M1 (H-GSP 98186). In the juveniles (H-GSP 5223 and 5224), the sutureof the maxilla in the orbit is horizontal and low. The zygomaticforms most of the anterior rim of the orbit, and contacts the lac-rimal superiorly. The anterior root of the zygomatic arch is robust,projecting laterally from the area over M2, and showing a broadsurface for the origin of the anterior masseter. Lacrimal exposureis small, both on the face and in the orbit and there is no lacrimaltubercle. A large lacrimal foramen is situated on the rim of theorbit, opening posteriorly. There is a strong postpalatine toruscontinued posteriorly into the broad pterygoid processes (H-GSP5224).

The postorbital process of the frontal forms the roof of theorbit. The posterior edge of the postorbital process bends stronglymedial and is continued into the temporal line (Fig. 3.2). Left andright temporal lines converge gradually toward the posterior partof the skull and only form a low sagittal crest in the posteriorpart of the skull in H-GSP 5224. In H-GSP 5223, the left andright temporal lines contact further anteriorly and the sagittal crestis longer. The fronto-parietal suture cannot be discerned in anyof the specimens. The roof of the interorbital region of H-GSP5223 is indented by seven shallow depressions arranged in a semi-circle. It is possible that these are puncture marks made by theanterior teeth of a predator or carrion feeder. Puncture marks werealso observed on the axis of a specimen of Diacodexis pakista-nensis from Barbora Banda (Thewissen and Hussain, 1990).

The squamosal makes up most of the lateral wall of the brain-case and is pierced by several emissary foramina. The squamosalroot of the zygomatic arch is broad, and the mandibular facet iswide and flat (H-GSP 5223). This facet is bounded posteriorly bya low postglenoid process that is narrower than the facet. Posteriorto the process is the external auditory meatus, and a small post-glenoid foramen occurs in the posterior aspect of the postglenoidprocess. The squamosal-alisphenoid contact extends antero-pos-teriorly on the medial side of the mandibular joint.

A process (alisphenoid tubercle) arises from the alisphenoidmedial to the mandibular joint (H-GSP 5223), and anteromedialto the postorbital process. A broad groove occurs between thealisphenoid process and the postglenoid process, possibly for themaxillary artery. Anterior to the alisphenoid process is the ovalforamen, from which a groove extends anteriorly. Medial to thisgroove is the base of the pterygoid process. The area medial tothe alisphenoid process and oval foramen is deeply excavated,and roofed by a thin layer of bone. This part of the alisphenoidprobably forms the anterior roof of the middle ear cavity.

The morphology of the lower jaw is known from an adult (H-GSP 5139) and a juvenile (H-GSP 5226). In the adult the hori-zontal ramus is slender and increases only slightly in depth fromanterior to posterior. Because the anterior of H-GSP 5139 is dam-aged, it is not possible to measure the length of the postcaninediastema in the adult, but the juvenile dentition shows a moderate(7 mm) postcanine diastema (Fig. 3.3). The symphysis terminatesbelow P1 in the adult (the anterior portion is missing) (Fig. 2.1)and beneath the diastema in the juvenile. In the adult the angleof the mandible extends behind the condyle, and the coronoid isdamaged but probably did not extend much above the condyle.

No upper incisors or alveoli are preserved in the type specimen,but a juvenile (H-GSP 5226) preserves an erupting small, spatu-late I3. C1 is a relatively large and labio-lingually compressed tusk.P1 and P2 are not known. P3 is three-rooted and triangular inshape. The metacone and paracone are high, equal in size andmoderately separated. The single lingual cusp (protocone) is con-ical and broader-based but slightly lower than the buccal cusps.A low preprotocrista descends from the protocone to the anterior

411MAAS ET AL.—NEW TAPIROMORPH FROM PAKISTAN

TABLE 1—Karagalax mamikhelensis n. gen. and sp., dental measurements (mm). Abbreviations: L—length; BL—buccal length; LL—lingual length; W—width; AW—anterior width; PW—posterior width.

H-GSP5142

H-GSP5143

H-GSP5144

H-GSP5145

H-GSP5139(r)

H-GSP5139

(l)

H-GSP5223

(l)

H-GSP5223(r)

H-GSP5224(r)

H-GSP5224

(l)

H-GSP5225

H-GSP5226(r)

H-GSP5226

(l)

H-GSP5229

H-GSP5230

H-GSP98186

(r)

H-GSP98186

(l)

P3 BLLLW

6.54.47.3

6.1——

P4 BLLLW

7.35.38.0

7.06.37.3

M1 LW

7.59.2

7.59.2

——

——

7.8—

7.89.2

8.38.7

8.28.4

M2 LW

——

9.29.7

——

8.48.7

8.48.7

M3 LAWPW

8.39.58.0

7.89.27.5

8.78.77.8

8.78.77.8

p1 LW

4.21.7

p2 LAWPW

6.02.53.2

p3 LAWPW

7.23.54.7

7.22.84.2

p4 LAWPW

7.54.34.8

7.53.84.8

m1 LAWPW

7.74.34.5

7.54.34.5

7.3–

4.7

———

10.05.65.6

m2 LAWPW

8.85.25.0

8.75.35.2

9.26.76.2

8.76.05.8

m3 LAWPW

12.55.85.0

125.55.0

10.7

DP1 LW

5.83.2

——

——

6.23.3

DP2 LW

6.1—

6.3—

——

——

DP3 LAWPW

6.6——

6.6——

———

———

DP4 LW

——

6.8—

——

7.26.8

base of the paracone and there is a very short crest (postproto-crista) that descends postero-buccally from the protocone but ter-minates before reaching the central basin. The preparacrista de-scends anterior to the paracone and terminates slightly lingual tothe base of the large parastyle. The anterior cingulum is narrowand the postcingulum is low and broad lingually. There is novestige of a posterolingual cusp. A sharp postmetacrista descendsposteriorly from the metacone and is continuous with a narrowbuccal cingulum, which terminates between the paracone andmetacone.

P4 is three-rooted and broader and longer than P3. The paraconeand metacone are high, subequal and moderately separated, as inP3. They are connected by a very short, straight centrocrista.There is a prominent parastyle anterobuccal to the preparacrista.The protocone is high, conical, and broader-based than the para-cone and metacone. It is located directly lingual to the paracone.A preprotocrista extends buccally from the protocone to the baseof the parastyle, where it intersects the preparacrista. There is avery small paraconule close to the buccal termination of the pre-protocrista. The postprotocrista descends buccally from the pro-tocone, deviates posteriorly at a small metaconule and terminatesat the base of the centrocrista between the paracone and metacone.The postcingulum is broad, with no evidence of a hypocone, and

rises buccally to converge with the postmetacrista. Buccal to themetacone the narrow cingulum bears a very low metastyle.

M1 has four prominent conical cusps. The metacone is slightlylower than the paracone and slightly displaced lingually, givingthe tooth a roughly trapezoidal outline. Both paracone and meta-cone are buccally convex. The preparacrista and postmetacristaare sharp. The centrocrista is straight and notched between theparacone and metacone. A prominent, laterally compressed par-astyle, about half the height of the paracone, rises from the an-terior cingulum anterior and slightly buccal to the paracone. Thetwo lingual cusps, protocone and hypocone, are conical, equal insize, and slightly lower than the buccal cusps. The protoloph ex-tends buccally and slightly anteriorly from the protocone and isconfluent with the preparacrista just posterior to the parastyle. Allspecimens have a small paraconule, nearly incorporated into theprotoloph but still distinct. The metaloph, which is slightly shorterthan the protoloph, extends buccally and slightly anteriorly fromthe protocone, then bends slightly posteriorly to join the centro-crista anterior to the metacone either high (H-GSP 5223) or closeto the base (H-GSP 5224). A small metaconule is variably pre-sent: it occurs on H-GSP 5223, on the left but not the right M1

of the type specimen and H-GSP 5224, and it is absent on theisolated anterior molars H-GSP 5142 and H-GSP 5143. There are

412 JOURNAL OF PALEONTOLOGY, V. 75, NO. 2, 2001

narrow anterior and posterior cingula but buccal and lingual cin-gula are very faint and in most cases incomplete.

M2 is virtually identical to M1, but slightly larger. A small par-aconule is present on both right and left M2 of all three specimens,and more prominent than in M1. A small metaconule is presenton H-GSP 5224 and H-GSP 5223 but not on the type specimen.

M3 is similar in morphology to M1 and M2 but with a lower,more buccally flattened metacone. The metacone is closer to theparacone than on M1 or M2, and the metaloph does not bendposteriorly as it reaches the ectoloph. There is a small paraconuleand metaconule on the type specimen, but the metaconule is ab-sent on H-GSP 5144 and H-GSP 5145. M3 is slightly smaller thanM2.

I2 and I3 are known only from slightly damaged erupting teeth(H-GSP 5226) but appear spatulate and somewhat procumbent.An erupting C1 (H-GSP 5226) is a labio-lingually compressedtusk, slightly smaller than the upper canine. It is separated fromthe premolars by a moderate diastema (H-GSP 5226). P1 is single-rooted. It has a single, high bucco-lingually compressed centralcusp with a short, slightly convex anterior crest and longerstraight posterior crest. The posterior crest forms the lingualboundary of a slightly broadened posterior basin, which is dividedbuccally by a deep groove. P2 is two-rooted with a high centralcusp (protoconid) sloping antero-buccally to a distinct, low para-conid, which is bordered buccally, and lingually by narrow cin-gula. A crest descends sharply in a postero-lingual direction fromthe protoconid and curves continuously around the lingual sideof a broad posterior basin (talonid). There is no entoconid. Thebuccal side of the posterior basin is bounded by a low crest thatrises posteriorly to a hypoconid. There is a buccal cingulum alongthe talonid.

P3 is two-rooted. A paracristid extends anteriorly from a highprotoconid and terminates in a low paraconid. There is a secondsmall cusp just lingual to the paraconid. A narrow anterior cin-gulid reaches from low on the buccal side of the tooth at the levelof the paraconid to the antero-lingual corner. The protocristid runspostero-lingually from the protoconid and terminates in a smallbut distinct metaconid. The cristid obliqua extends from the centerof the posterior trigonid wall postero-buccally to the hypoconid.A buccal cingulid reaches from the posterior base of the hypo-conid anteriorly to the base of the posterior trigonid wall. Thetalonid basin is bounded buccally by a high hypoconid but thelingual border is low with no distinct entoconid.

P4 is more molariform than P3 and nearly rectangular in outline,with a high trigonid and lower talonid. The protoconid is slightlyhigher than the metaconid; the two cusps are joined by a trans-verse, notched protolophid. A low paracristid curves anteriorlyand lingually from the protoconid, but there is no paraconid. Thecristid obliqua extends anteriorly and slightly lingually from thehypoconid down to the protolophid just buccal to the midline.The hypoconid is low and cuspate. The low hypocristid extendslingually a short distance from the hypoconid. There is a smallentoconid on the right P4 of the type specimen but not the left. Anarrow posterior cingulum bears a small hypoconulid.

M1 is nearly rectangular and the trigonid is very slightly nar-rower than the talonid. The talonid is lower than the trigonid.Protoconid and metaconid are conical and equal in size and joinedposteriorly by a notched, transverse protolophid. There is no me-tastylid and the metaconid is not twinned (Hooker (1994) treatsthese as separate features). A low paracristid runs anteriorly fromthe protoconid, then turns sharply lingual along the anterior toothborder. There is no paraconid. A narrow cingulid extends fromthe antero-lingual corner of the tooth around the buccal side andin some specimens is continuous with a postcingulid. The cristidobliqua extends anteriorly and slightly lingually down from thehypoconid to the base of the protolophid below the protoconid.

Hypoconid and entoconid are high and conical, equal in size, andjoined by a transverse, notched hypolophid. There is a small, lowhypoconulid located posterior to the hypolophid on the postcin-gulid, midway between the hypoconid and entoconid. The hy-poconulid is connected to the hypolophid below the notch by avery short, low crest. M2 is virtually identical to M1, but slightlylarger, and with the trigonid very slightly wider than the talonid.The buccal cingulid is slightly broader than in M1. M3 is similarin morphology to M1 and M2, but the postcingulid is expandedas a large, broad hypoconulid lobe, which tapers lingually. Thecrest that forms the posterior border of the hypoconulid lobe bearsa variable number of small cuspules. It is connected to the baseof the hypolophid by a low crest, just buccal of midline.

The deciduous upper dentition is best preserved in H-GSP5224, and the lower dentition in H-GSP 5226. DP1 is narrow,elongate and two-rooted with a single large central cusp. A longridge descends posteriorly from the cusp to a posterior cingulum,and a shorter ridge descends anteriorly to a short anterior cingu-lum bearing a small parastyle. The cingulum extends along theposterior two-thirds and is especially wide posteriorly. DP2 is tri-angular with a long, oblique lingual edge. The paracone is thelargest cusp. The metacone is distinct, but smaller and close tothe paracone. The lingual half of the tooth appears to be an ex-panded shelf with a single, laterally compressed cusp on the cin-gulum. DP3 is subquadrate; it is narrower anteriorly and the lin-gual edge is oblique. The paracone and metacone are roughlyequal in size (the latter may be slightly larger) and they are linkedby a notched centrocrista. The hypocone is larger than the pro-tocone. Both are conical, and smaller than the buccal cusps. Thereis a prominent parastyle. The metaloph extends buccally and an-teriorly from the hypocone to the antero-lingual base of the meta-cone, just posterior to the centrocrista notch and bears a smallmetaconule. The protoloph is short and low, and terminates lowon the ectoloph anterior to the paracone. DP4 is quadrate andmolariform. It is morphologically similar to M1, but slightly nar-rower.

The single-rooted tooth anterior to DP2 in H-GSP 5226 is iden-tical to P1 (H-GSP 5139) and may or may not be deciduous. DP2

is two-rooted and has a badly damaged crown. It appears to havea short, broad talonid, but no cristid obliqua. DP3 is long andsubmolariform. A high paracristid extends buccally and then pos-teriorly from the antero-lingual paraconid. The protoconid islarge, and a crest curves posteriorly and lingually to a lower meta-conid, which is about the same height as the paracristid. Thecristid obliqua extends anteriorly from a prominent hypoconid.The talonid is broad and basined. The entoconid is slightly smallerthan the hypoconid and the hypocristid was probably small orabsent. DP4 is fully molariform. It is very similar to M1 but slight-ly smaller, and the trigonid is slightly broader than the talonid (H-GSP 5226).

A number of postcranial elements were preserved at the typelocality though none was anatomically associated with Karagalaxdental material. However, these bones compare well with tapiroidpostcranials (e.g., Heptodon, Radinsky, 1965a), and are consistentwith the tooth size of Karagalax. The fauna contains no othertaxa of this size and we have referred them to Karagalax. As withthe other fossils from this locality, most postcranial elements havebeen deformed post-mortem to various degrees.

No humerus is preserved completely, and the most intact spec-imen is a juvenile lacking the proximal epiphysis (H-GSP 98179).It is 79 mm long. The humeral head of this specimen is deformedbut was probably longer antero-posteriorly than the humeral headof Heptodon (Radinsky, 1965a). The deltopectoral crest is nearlyinvisible. The olecranon fossa is wide and deep and the capitulumis narrow. The epicondyles are weak. There is no supracondylarforamen.

413MAAS ET AL.—NEW TAPIROMORPH FROM PAKISTAN

Only distal parts of the radius are preserved. A radial epiphysis(H-GSP 98182) shows a deep groove for extensor carpi radialis,and medially a well-developed styloid process. This process is onthe edge of the facet for the scaphoid, which is slightly concaveanteriorly and convex posteriorly. The facet for the lunar is con-cave anteroposteriorly but more or less square in outline andsmaller than the facet for the scaphoid.

A proximal ulna (H-GSP 98183) lacks the tip of the olecranon.Its humeral facet is narrow and ends proximally in a prominentprocess. The distal part of the semilunar notch is expanded for abroad articulation with the radius, both medial and lateral to thesemilunar notch. The shaft of the ulna distal to this tapers stronglyand at midshaft the bone is much deeper (anteroposteriorly) thanwide (mediolaterally).

Only three juvenile, incomplete femora are preserved. Theyshow a large, flattened third trochanter at the base of a broadtrochanteric fossa (H-GSP 98167). The third trochanter is exten-sive proximo-distally. The shaft is gracile (H-GSP 98161). Thetibia has a strong and wide tibial crest, and its condyles are strong-ly convex anteroposteriorly (H-GSP 98164). The distal tibia hasa strong central crest, implying that Karagalax has a deeplygrooved astragalus (H-GSP 98184).

The astragalus is deformed by antero-posterior compression (H-GSP 98172). It shows a deeply grooved trochlea, and has a longneck. The concavity of the ectal facet is oriented more or lessobliquely to the axis of the bone. The sustentacular facet is small.A preserved fragment of a distal metapodial (H-GSP 98171) hasa head that is strongly convex in anteroposterior and mediolateraldirections. A crest occurs on the posterior part of the head (H-GSP 98178). It is placed asymmetrically on the head implyingthat this is a lateral metapodial. The distal phalanx (H-GSP98177) is similar to Heptodon, but is more expanded distally. Itis 11 mm in length and 8.5 mm in distal width.

COMPARATIVE MORPHOLOGY

The affinities of Karagalax lie unquestionably with the basaltapiromorphs. Karagalax differs from primitive hippomorphs inthe termination of the molar metaloph relatively high on the ec-toloph and reduction of the post-canine diastema, both tapiro-morph synapomorphies (Hooker, 1989). It likewise shares thethree tapiromorph synapomorphies identified by Froehlich (1999):lophoid molar metaloph, angle of convergence of upper molarbuccal and lingual cusp less than 908, and crenulated M3 lobe.Within Tapiromorpha, Karagalax most resembles the isectolo-phids. Like all isectolophids (Radinsky, 1963; Ting, 1998) Kar-agalax has molars with straight pre- and postmetacristae and, likeall isectolophids except the lophodont Isectolophus, it retains theprimitive tapiromorph features of small but distinct molar conules(Fig. 1.1) and notched lower molar cross-lophs (Fig. 2.1). Amongisectolophids Karagalax is more derived than Cymbalophus. Itsmolars are more lophodont, bearing a stronger metaloph, a lessnotched hypolophid, and lacking a metastylid [twinned metaconidof Froehlich (1999) and Hooker (1994)]. Karagalax differs fromCardiolophus and North American Homogalax in a combinationof primitive and derived features of the cheek teeth, as well itssmaller size. In contrast to most Homogalax, P3 of Karagalax hasa short postprotocrista, the P4 metaconid is lingual rather thanposterolingual to the protoconid, and DP2 has a lingual cusp. Allof these are more derived features. On the other hand, its slightlyless lophodont molars (more deeply notched protolophid and low-er, more deeply notched hypolophid) are more primitive thanNorth American Homogalax and Cardiolophus, and it certainlyhas not reached the degree of lophodonty or premolar molariza-tion of Isectolophus (Radinsky, 1963). In contrast to Isectolophus,Karagalax’s molar conules are not completely incorporated into

cross-lophs, the protolophid and hypolophid are notched, and P3

does not have a hypocone.Comparisons with the Asian isectolophids Orientolophus and

Homogalax wutuensis are based on published descriptions (Ting,1993). Both are more primitive (less lophodont) than Karagalax.Orientolophus, which is slightly smaller than Karagalax, retainsmetastylids on all three lower molars, has more deeply notchedprotolophids and hypolophids, and has more distinct paraconulesand metaconules. Homogalax wutuensis, though more lophodontthan Orientolophus (Ting, 1993), retains a large molar metastylid,which is absent in Karagalax. The unnamed isectolophid fromthe Wutu Basin has not been described in detail, but is reportedto have large metastylids, metaconules, and paraconules (Tongand Wang, 1998), all of which distinguish it from Karagalax.

Karagalax resembles the problematic Indian Sastrilophus onlyin the primitive retention of small molar conules. Karagalax hasa more molariform P3 than Sastrilophus (paracone and metaconebetter differentiated and the metacone equal height to paracone),and shares none of the diagnostic generic characters of the genus[flattened metacones, which are significantly lower than the para-cones, large parastyles, and upper molars that are less transversethan those of North American Homogalax, Isectolophus, or theAsian deperetellids (Sahni and Khare, 1971)]. The most strikingdifference between Karagalax and Sastrilophus is the more buc-cal position of the metacone in Sastrilophus, which results in anearly square outline of M1 and M2 (Fig. 4.2). This contrasts withthe typical tapiromorph trapezoidal molar perimeter and distin-guishes Sastrilophus from all other tapiromorphs, including Kar-agalax.

The two juvenile skulls of Karagalax are the only cranial fos-sils yet described for early Tertiary Asian tapiromorphs, with theexception of the recently reported skull of the late Eocene lophi-aletid, Eoletes (Lucas et al., 1997). The skulls are characterizedby a suite of primitive tapiromorph (isectolophid) features and arebroadly comparable to Cardiolophus (Gingerich, 1991). Featuresshared with Cardiolophus include a postorbital process that formsa broad roof over the orbit, absence of a lacrimal tubercle (seeCardiolophus UM 78915), and presence of an alisphenoid tuber-cle (see Gingerich, 1991, fig. 8). Both Karagalax and Cardiolo-phus differ from more derived tapiromorphs such as the middleEocene helaletid Heptodon (Radinsky, 1965b) in their wider glen-oid fossa, absence of a lacrimal tubercle, and in the position ofthe posterior opening of the alisphenoid canal lateral rather thanposterior to the pterygoid process. Like Cardiolophus and Hom-ogalax, Karagalax has a moderately shortened postcanine diaste-ma, a feature that Hooker (1989) interpreted as an isectolophidsynapomorphy. However, Systemodon and Cymbalophus [whichHooker (1989, 1994) considered hippomorphs] have moderatelylong postcanine diastemata (Froehlich, 1999), and some lophiale-tids also have short postcanine diastemata (Hooker, 1989; Lucaset al., 1997). Based on the degree of nasal incision, none of theisectolophids, including Karagalax, appear to have had a probos-cis, but the nasal incision is deeper in Karagalax than in Homo-galax, and similar to that of Heptodon, where it reaches P1 insome specimens (Radinsky, 1963).

Other than shared primitive tapiromorph dental features, Kar-agalax shows no resemblance to the more derived Asian tapiro-morph families, Deperetellidae, Helaletidae, and Lophialetidae. Itlacks the diagnostic deperetellid characters of molarized premo-lars, high-crowned, bilophodont molars, reduced or absent M3

metacone, reduced paralophid and cristid obliqua, and absence ofM3 hypoconulid (Radinsky, 1965b; Hooker, 1989). It lacks diag-nostic helaletid characters such as long post-canine diastema, flat-tened and shortened M1–2 metacone, reduced M3 metacone, andreduction/absence of the lower molar cristid obliqua (Radinsky,1965b; Hooker, 1989). Karagalax shows some similarities to the

414 JOURNAL OF PALEONTOLOGY, V. 75, NO. 2, 2001

FIGURE 4—1, Karagalax mamikhelensis, H-GSP 98186 left P4-M3. 2,Sastrilophus dehmi, LUVP 15001, left M1–3. 3, Chasmotherium mcken-nai, LUVP 15011, left M1–3, referred to Kalakotia simplicidentata. 4,Homogalax sp., USGS 19428, left P4-M3. 5, Cardiolophus radinskyi,UM 78915, left P4—M3. Not to scale.

primitive lophialetid Kalakotia, but is clearly more primitive. Kar-agalax and Kalakotia both lack molar metastylids and have morelophodont protolophids and hypolophids than Orientolophus, butthe trend towards molar lophodonty is much more pronounced inKalakotia than Karagalax. Thus, upper molars of Karagalax re-tain weak metaconules and paraconules and have buccally con-vex, less lingually deflected metacones than Kalakotia (compareFig. 4.1, 4.3, especially M3). Likewise, the deciduous premolarsof Karagalax are less molariform—they are narrower and longerthan those of Kalakotia (LUVP 15011, type of Chasmotheriummckennai) and the DP3 paracone and metacone are closer together.In addition to these more primitive features, Karagalax lacks anyof the diagnostic generic characters of Kalakotia (long postcanine

diastema, molars with convex paracones and slightly concavemetacones, and a V-shaped protoloph-metaloph loop on P4). Kar-agalax differs from lophialetids other than Kalakotia, in the ab-sence of a complete U-shaped lophoid loop (formed by pre- andpostprotocristae and an antero-posteriorly lengthened protocone)on P3–4 (Hooker, 1989). It has a degree of nasal incisure similarto that of the middle Eocene lophialetid Eoletes, but lacks itsknob-like postorbital process and typical lophialetid dental fea-tures (Lucas et al., 1997, fig. 3).

The postcrania of Karagalax show a combination of features,some of which are more primitive and some more derived thanother isectolophids, but none of the derived features suggest acloser relationship to any of the more advanced tapiromorph fam-ilies. Primitive characters include the long neck of the astragalus(Fig. 3.4) [longer than that of Hyracotherium, (AMNH 48663)]and the comparatively large third trochanter, both of which areless advanced than homologous features of Homogalax (Rose,1996). However other features are more advanced. These includeweaker humeral epicondyles, deeper olecranon fossa, and a moregracile femur. These features are even more derived than in thehelaletid Heptodon (Radinsky, 1965a), and suggest that Karaga-lax was more cursorial than the North American isectolophids.

PHYLOGENY

The recent phylogenetic analysis of basal perissodactyls byFroehlich (1996, 1999) allows us to summarize the relationshipsof Karagalax within the broad context of tapiromorph phylogeny.We scored Karagalax and Kalakotia (ONG/K/10 and LUVP15011, holotype of Chasmotherium mckennai) according toFroehlich’s character set (see Appendix). We determined the mostparsimonious placement of Karagalax and Kalakotia on Froeh-lich’s cladogram (Froehlich, 1999, fig. 5) by exploring a numberof combinations, using MacClade 3.0 (Maddison and Maddison,1992). The results are illustrated in Figure 5. The phylogeneticposition of Karagalax reflects the suite of primitive and derivedfeatures that characterize its dentition. Our analysis shows thatKaragalax is derived relative to Orientolophus and Systemodon/Cymbalophus but primitive relative to Cardiolophus and NorthAmerican Homogalax. Kalakotia appears as sister taxon to Kar-agalax, and both are less derived than Cardiolophus and Homo-galax.

We caution that these phylogenetic assessments are provisional,especially with respect to the Asian tapiromorphs. There are sev-eral reasons for this. 1) We did not re-run Froehlich’s charactermatrix with the added Indo-Pakistani taxa because we did nothave access to all of the taxa included in his analysis, and becausethe additional missing data for the Indo-Pakistani taxa might de-stabilize his original tree. 2) Some inconsistencies may be intro-duced by adding independently scored taxa to an existing matrix.3) Some important basal tapiromorphs were excluded—Sastrilo-phus was not included because so few characters can be scoredfor that taxon, and Homogalax wutuensis material was not avail-able for scoring. 4) The more derived Asian tapiromorphs (e.g.,lophialetids) are not included—their omission undoubtedly weak-ens our assessment of the phyletic position of Kalakotia. Never-theless, the phylogenetic relationships outlined here represent aworking hypothesis, which will be tested by future finds from theearly Tertiary of Asia.

The finding that Karagalax is more derived than the ChineseOrientolophus suggests that tapiromorphs, which were alreadydistinct from hippomorph perissodactyls (Froehlich, 1999), dis-persed to the Indian subcontinent from the Asian continent. Al-though the phylogenetic position of Karagalax does not directlyrefute the possibility that perissodactyls originated on the Indiansubcontinent and dispersed to Asia (Krause and Maas, 1990), it

415MAAS ET AL.—NEW TAPIROMORPH FROM PAKISTAN

FIGURE 5—Cladogram of basal tapiromorph genera from Froehlich(1999, fig. 5), with the addition of Karagalax mamikhelensis and Ka-lakotia simplicidentata. See text for explanation of cladogram construc-tion and Appendix for character state scores for Karagalax and Kalak-otia. Unequivocal synapomorphies for internode A include Froehlich’scharacters 36(1) P4 postprotocrista lophoid, joining ectoloph (reversedin Cardiolophus), 53(2)—upper molar protolophs sub-lophoid, 78(1)—P4 slightly smaller than M1, 102(1)—M3 heel long, 100(3)—M3 hy-poconulid basin large (except Kalakotia), and 106(3)—wear patternvertical with elaboration along facets 2, 2a, 8, 7, 7a. Unequivocal syn-apomorphies for internode B include Froehlich’s characters 40(1) uppermolar parastyle buccal to line connecting paracone and metacone and64(1)—upper molar much wider than long. Internode C is supportedby Froehlich’s characters 66(1)—upper molar protocone-hypocone sep-aration greater than paracone-metacone separation, and 72(1) P3 hy-poconid large.

is most consistent with the more broadly accepted idea that pe-rissodactyls first appeared in continental Asia and dispersed fromthere to other areas, including Indo-Pakistan (see Beard, 1998).Preliminary biogeographic assessments of an as yet undescribedmammal fauna from the upper Ghazij Formation in southern Pak-istan also appear to support this view (Gingerich et al., 1999).

AGE OF BARBORA BANDA FAUNA

The Eocene mammalian fauna from Indo-Pakistan includesboth a diverse endemic component and a number of specieswhose affinities could lie with Holarctic forms (Sahni and Kumar,1974; West, 1980; Lucas and Schoch, 1981; Sahni et al., 1981;Kumar and Jolly, 1986; Thewissen et al., 1987; Gingerich, 1990;Krause and Maas, 1990). There is a growing consensus that, by

the early Eocene, the Indian subcontinent was in contact with theAsian plate (Curray et al., 1982; Wells, 1983; Besse et al., 1984;Patriat and Achache, 1984; Thewissen, 1990; Thewissen and Mc-Kenna, 1992), although an epicontinental sea may have separatedthe land masses. This suggests that faunal exchange could haveoccurred near the Paleocene-Eocene boundary. However, until re-cently the best known Eocene Indo-Pakistani faunas have beenthose middle and late Eocene in age and, for the most part, havebeen dominated by endemic taxa. H-GSP Locality 300 appears tobe an exception.

Geological studies have documented the distinctiveness of theMami Khel Formation, which occurs west of the Indus River,from the Kuldana Formation, exposed largely east of the Indus(Meissner, 1974; Pivnik and Wells, 1996). Several lines of bio-stratigraphic evidence suggest that the three Mami Khel localities(H-GSP Locality 300 and two other nearby localities, H-GSP Lo-calities 9710 and 9712), which together comprise the Banda DaudShah fauna (Thewissen et al., in review), are older than the Kul-dana Formation localities, and probably of early Eocene age.First, the primitive dichobunid artiodactyl Diacodexis pakistanen-sis, which is the most common species at H-GSP Locality 300,is generally considered to be at the base at the artiodactyl radia-tion (Gentry and Hooker, 1988; Theodor, 1996). Second, two ofthe rodent specimens from Barbora Banda locality are thought torepresent species that predate the chappatamyid radiation (deBruijn et al., 1982). Third, plesiadapoids and arctocyonids, groupsthat are most abundant in Paleocene faunas of other continentsand decline in diversity in the early Eocene, occur in the BandaDaud Shah area (H-GSP 9710 and H-GSP 9712) (Thewissen etal., in review). Fourth, all of these faunal elements are absent fromearly middle Eocene localities to the east, which are dominatedby more derived endemic Asian ungulates and rodents (Russelland Zhai, 1987; Leinders et al., 1999; Thewissen et al., in review).The evolutionary position of Karagalax, as the most primitiveperissodactyl yet described from Indo-Pakistan, provides a fifthline of evidence to support the conclusion that H-GSP Locality300 and the Banda Daud Shah fauna, is early Eocene in age.

ACKNOWLEDGMENTS

This paper is a contribution to the Howard University–Geolog-ical Survey of Pakistan project ‘‘Cenozoic Stratigraphy and Mam-malian Evolution in Pakistan’’. We thank those involved since itsinception in 1974 for their contributions and support. We are es-pecially indebted to the Director General, Geological Survey ofPakistan and his staff, who have given us their fullest cooperationand support at all stages of the work. We are particularly gratefulto M. Arif, H. de Bruijn, I. Cheema, N. Khan, A. J. van derMeulen, J. Munthe, S. -D. Nio, and R. M. West for undertakingthe fieldwork and for collecting the specimens. Thanks to D.Froehlich and an anonymous reviewer for helpful comments onan earlier draft of this manuscript. Support was provided from theSmithsonian Institution, National Science Foundation, and Na-tional Geographic Society to S. T. H., and National GeographicSociety (5536-95) and National Science Foundation (currentlyEAR-9902830) to J. G. M. T. and S. T. H. The staff of the Instituteof Earth Sciences, University of Utrecht and E. M. Williams,NEOUCOM, prepared the specimens.

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APPENDIX.—Character scores for Karagalax mamikhelensis and Kalakotiasimplicidentata for character matrix of Froehlich (1996, 1999).

Karagalax10?000??1? 0001102010 ??10000-10 0100010110 1111100030 01212210010000201010 00202201130 00-20411012 30101213011 203?????01????200101??

Kalakotia??????????? ??????????? ??????????? ?????201010 00200-0012 02300021001201010002 0000113000 2203110223 21011130102 03???????????????????

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