Geol. Bull. Univ. Peshawar, Vol. 36, pp. 73-83, 2003

M. SADIQ MALKANI Paleontology and Stratigraphy Branch, Geological Survey of Pakistan, Sariab Road, Quetta, Pakistan

ABSTRACT: First Kimmeridgian (Late Jurassic) dinosaur fossils have been discovered in the lowest horizon of Sembar Formation just near the contact of ZidiKhiltan Limestone, in the Sun Chaku locality of Karkh area, and Lakha Pir Charo locality of Zidi area, Khuzdar district, Balochistan, Pakistan. Fossils from the lowest part of the Semba? Formation in Kirthar Range, Pakistan, provide the first remains of Late Jurassic Titanosaurian dinosaur from Pakistan. However, the specimens collected to date include the poorly preserved, fragmentary and incomplete cross section of femur, metatarsals/metacarpals, ribs and proximal portion of fibula and distal portions of neural spines/diapophysis/zygapophysis. To acknowledge the tribe and locality, Brohisaurus kirthari, a new genus and species of Titanosauria, is erected on the basis of characters observed which are elliptical/eccentric nature of femoral cross section, well developed of proximal scar on proximal fibula, possible pneumatic cavities in the anterior ribs, massiveness in the distal ribs, less arced D shape cross section of rib and some resemblance of proximal fibula and ribs cross section to the Late Cretaceous Pakistani Titanosaurs. The degree of elliptical nature is diflerent from Late Cretaceous dinosaur fauna of Pakistan.

This discovery of Jurassic dinosaur fossils from Pakistan is expected to develope great interest in local and global scientzjic community which will open new avenues of research such as exploration of any age of Mesozoic dinosaurs and their phylogeny and paleobiogeography. Dinosaurs were domrmnantland animals during most of the Mesozoic era from 225 to 65 million years ago, but became extinct at its end. Thick piles of Mesozoic sedimentary rocks are exposed in many parts of Pakistan and it is encouraging for the exploration of every age of Mesozoic dinosaurs.

Indo-Pakistan initially interlocked with the Gondwana landmasses of Africa, South America, Australia, Antarctica and Madagascar early in the Mesozoic, drifted northward during the Cretaceous to collide with Laurasian landmasses during the Cenozioc. Indo-Pakistan thus appears to have experienced a 100-million-year period of isolation during a 9,000 kilometer migration across the equator that can be expected to have influenced the character of its native biota. Fossils from this period of biogeographic fsolation are relatively scarce. So far, the Late Cretaceous (Maastrichtian)Lameta Formation of India has sewed as the sole source qf information on Cretaceous vertebrates of the Indo-Pakistan subcontinent. But new discoveries including a variety of large vertebrate that indicate five new genus and species of herbivorous Titanosaurian Sauropod and one new genus and species of carnivorous Abelisaurid Theropod Dinosaurs and Baurosuchid Pabwehshi Pakistanensis (Mesoeucrocoreptilia) fossils. Zkse discoveries of Jurassic and Cretaceous Arhosaurian reptiles from Pakistan act as a aik~tonefor assessing paleobiogeography and phylogeny.

INTRODUCTION great interest in local and global scientific community which will open new avenues of First Kirnrneridgian (Late Jurassic) dinosaur research such as paleobiogeography and fossils have been discovered first time in phylogeny. A more precise taxonomy, Pakistan in the lowest horizon of Sembar anatomy, phylogeny, skeletal variation, track Formation just near the contact of ways evolution, etc., of dinosaurs will be Zidi/Chiltan Limestone, in the Sun Chaku clear after articulated collections. Dinosaurs locality (coordinates; 27'50' 40" N; 67' 09' were giant reptiles, these animals were quite 66" E) of Karkh area, and Lakha Pir Charo anomalous from normal evolutionary pattern locality (coordinates; 27' 37' 35" N; 67" 02' of other species and genera and so big in size 10" E), of Zidi area, Khuzdar district, that they are of great interest for taxonomists, Balochistan, Pakistan (Fig. 1). The Sun taphonomists, vertebrate paleontologists and Chaku locality is 15 kilometers from Karkh ecologists. Dinosaurs were dominant land village and 85 kilometres east of Khuzdar. It animals during most of the Mesozoic era from is accessible through fair-weather road. Charo 225 to 65 million years ago but became locality is 15 kilometers south of Bhalok extinct 'at its end. One of the unresolved village and 60 kilometres east of Khuzdar. mysteries of geological sciences is the existence of world's largest reptiles the Late Jurassic/Kimrneridgian dinosaur dinosaurs on planet earth during the periods fossils occur in the lowest horizon of Sembar of Triassic to Cretaceous and then their Formation and comprised dominantly shale abrupt extinction at the end of Cretaceous, which is enriched by iron nodules in both Thick piles of Mesozoic sedimentary rocks localities and have a light yellow color in the are exposed in many parts of Pakistan and the Charo area. Jurassic dinosaur fossils are Cretaceous-Tertiary boundary is well exposed found as fragmentary in the residual at many localities and it is encouraging for colluviums and bed rock dipping about 30 the exploration of every age of Mesozoic degree east at Sun Chaku and 25 degree east dinosaurs and reIated problems like at Lakha Pir Charo Iocality. Present study is extinction, taxonomy, paleobiogegraphy and based upon surface collections. Fossils from phylogeny. the lowest part of the Sernbar Formation in Kirthar Range, provide the first remains of Indo-Pakistan initially interloked with the the Late Jurassic Titanosauria from the Gondwana landmasses of Africa, South Pakistan. However the specimens collected to America, Australia, Antarctica and date include the poorly preserved, Madagascar early in the Mesozoic, drifted fragmentary and incomplete cross section of northward during the Cretaceous to collide femur, metatarsaIs/metacarpals, ribs and with Laurasian landmasses during the proximal portion of fibula and distal portions Cenozioc. Indo-Pakistan thus appears to have of neural spines/diapophysis/zygapophysis. experienced a 100 million year period of isolation during a 9,000 kilometer migration This discovery of Jurassic dinosaur across the equator that can be expected to fossils from Pakistan is expected to develope have influenced the character of its native biota. Fossils from this period of new genus and species of carnivorous biogeographic isolation are relatively scarce. Abelisaurid Theropod Dinosaurs and ~husfar, the late Cretaceous (Maastrichtian) Baurosuchid Pabwehshi pakistanensis Lameta formation of India has served as the (Mesoeucrocoreptilia) (Wilson et al., 200 1) sole source of information on Cretaceous fossils. These discoveries of Jurassic and vertebrates of the Indo-Pakistan subcontinent Cretaceous Arhosaurian reptiles from (Wilson, et al., 2001). But new discoveries Pakistan provide a vantage point and act as a including a variety of large vertebrate that milestone for for the entering of Indo- indicate five new genus and species of Pakistan subcontinent into new hypothesis of herbivorous Titanosaurian Sauropod and one Paleobiogeography and Phylogeny.

AShahdad kot Fig. 1. Index map showing the Sun Chaku and Charoh localities, Kirthar Range, Pakistan. GENERAL GEOLOGY Paleocene Dungan Formation, Eocene Shaheed GhatILaki (Ghazij group) and Sun Chakku and Charoh are located in the Kirthar Formations and Oligocene Nari and central part of Kirthar Range of Balochistan Gaj Formations and Miocene-Pliocene province, Pakistan. Sedimentary rocks Manchar Formation (Table 1) . ranging in age from Jurassic to Oligocene along with Quaternary surficial deposits occur The Quaternary deposits have been in the area. Sedimentary bed rocks such as further subdivided in to Pleistocene Dada limestone, shale, marl and minor sandstone Formation: Such as recent Terrace gravel bodies are divided in to several lithologic deposits, fan gravel deposits, colluviurns units, which in ascending order are Jurassic deposits, sand, silt and clay deposits (non Zidi Formation and Chiltan Limestone, cultivated and cultivated lands) and modern Jurassic-Cretaceous Sembar Formation, channel deposits. Cretaceous Gom, Parh, Mughallcot, Fort Munro and Pab Formations, Cretaceous- Stratigraphic sequence of the area is Paleocene Khadro and Rakhi Gaj Formations, tabuladd as follows:

TABLE 1. STRATIGRAPHIC SEQUENCE OF AREA

Age Formation Lithology Q Recent Modern channel deposits Sand, Gravel, sand, silt and clay silt and clay deposits Sand, silt and clay with minor gravel (cultivated lands)

and Sand, silt and clay deposits Sand, silt and clay with minor gravel (non-cultivated lands) CoIluvium deposits Boulder, pebbles, cobbles, with sand silt and clay. Fan gravel deposits Poorly consolidated gravel, sand, silt and clay. Sub-Recent Terrace gravel deposits Poorly consolidated gravel, sand, silt and clay. Angular Unconformity Pleistocene Dada formation Calcareous white clays. Angular Unconformity Gaj Formation Limestone, shale and sandstone Oligocene Nari Formation Sandstone, limestone and shale.

Disconformity

Kirthar Formation Limestone, marl and shale. Eocene Shaeed GhaWFomtion Mainly shale with minor marl / (Ghazij Group) limestone and mudstone / siltstone. Paleocene Dungan Formation Limestone, mat1 and shale Disconformity Rakhi Gaj Formation Shale, mudstone, siltstone, sandstone & limestone Khadro Formation Limestone with minor shale and sandstone Late Pab Formation Sandstone with subordinate shale Fort Munro Formation Limestone, shale and coquina beds Mughal Kot Formation Shale with minor marl/limestone and coquina beds

Early Parh Formation Limestone with minor marl and shale

Goru Formation Shale and marl with minor limestone

Sembar Formation Mainly shale with minor marl and mudstone Late Disconfonnity

Chiltan Formation Limestone with insignificant shale ZidiIShirinab Formation Limestone, shale, marl and minor sandstone

ANATOMY AND POSSIBLE 96-K, MSM-97-K, MSM-98-K, MSM-101 -K, PHYLOGENY MSM-102-K, MSM-103-K, MSM-104-K, MSM- 105-K, and MSM- 106-K. Femoral, Present findings consist of fragments of ribs, metatarsals and metacarpals cross appendicular skeleton such as femoral shaft sections, broken neural arch laminaelneural cross section (Fig. 2, MSM-86-k), cross spine and proximal fibula (Figs. 2 & 3) have section of metatarsals and metacarpals, been discovered during this study. The type proximal part of fibula (Fig. 3, MSM-88-K), specimens are part of collection of and fragments of axial skeleton such as pieces fragmentary pieces, made in Sun Chaku locality of ribs (Fig. 3, MSM-87-k, MSM-92-k, on slope of small hill in the clay rich rocks. MSM-93-k, MSM-94-k) and possible neural Referred specimens: MSM-95-K, One spineslneural arch laminae (MSM-95-k, broken piece of neural arch laminae was MSM-96-k). Only one piece of bone (MSM- found from the clay rich horizon of Lakha Pir 96-K) is found in the Charoh locality and all Charo area. other pieces of bones were found in the Sun Horizon and locality: Lowest clayhhale Chakku locality. The exposures of this horizon horizon of Sembar Formation just at and near are wide but upto now not followed for the contact of ZidiJChiltan limestone and detailed sampling. Sembar Formation, Sun Chakko Locality, Karkh area, Khuzdar district, Baluchistan, Systematic Paleontology (Taxanomy) Pakistan. Dinosauria Owen, 1842 Age: Late Jurassic (Kimmeridgian) Order Saurischia Seeley ,1888 Diagnosis: All the holotypic and referred Suborder Sauropodomorpha Huene, 1932 specimens are weathered, fragmentary, Infraorder Sauropoda Marsh, 1878 poorly preserved and replaced by ferruginous Family Titanosauridae Lydekker ,1 885 minerals. However, the ellipticity of femoral New Genus and Species, Brohisaurw Kirthari cross section, well-developed proximal scar on proximal fibula, simple slightly convexo- Holotype: MSM-86-K, MSM-87-K, MSM- convex (biconvex) ribs; and pneumatic 88-K, MSM-89-K, MSM-90-K, MSM-91-K, cavities in the anterior ribs are the best MSM-92-K, MSM-93-K, MSM-94-K, MSM- character to assign them Titanosaurs. The new genus and species names are purely known by the greater width and pneumatic tentative. nature of ribs. One piece shows the distal ends Etymology: New Genus and Species name and it is weathered and replaced by iron and Brohi is derived from the Brohi tribe of the mud but one piece seems to be massive. area, saurus means reptile and the species specific epithet represents the Kirthar range Both the central rib cross section are which is the host of Jurassic dinosaur. weathered and fragmentary but show some indication of pneumatic cavity which is a DESCRIPTION characteristics of Titanosauria. Most of the fragments are nearly flat and straight and Appendicular Skeleton none show strong curvature. The shape of transverse section varies in different ribs and Femoral cross-section: Broken femoral cross in different parts of the same ribs, from section (Fig. 2, MSM-86-K) having medio- narrowly elliptical to plano-convex. The lateral width of 12 crn and antero-posterior longest fragments measures 12 cm in length depth of 5.5 cm but its one side is weathered so with a breadth of the proximal end of 8.5 cm it may be of 6 cm. So this eccentric ellipticity and distal end of 7.5 cm. The widest fragment (12 x 6 cm) recalls the specimens belong to the is 8.5 cm. The width of ribs cross-section vary Titanosaurs. Femoral cross section is not from 8.5 to 7.5 cm and depth vary from 3 to 2.5 hollowed. Three smaller pieces of broken nature cm within a broben piece of 12 cm long. may belong to femora/humerus are shown in Fig. 1 (MSM-89-K, MSM-91-K, MSM-102-K). Neural spine fdiapophysial laminae: Two broken pieces (Fig. 3; MSM-95-K, MSM-96- Proximal Fibula: Proximal portion of fibula K) may belong to neural spine or diapophysial (Fig. 3, MSM-88-K) was found in Sun lamina. MSM-95-K is curved and it has Chakko locality. Its width is 10 cm and depth elongated lineation or grooves/rugosities for is 4 cm. Its one side form a well-developed the attachment of muscles. scar for attachment of proximal tibia. The fibular scar also recalls that the specimen The entry of Pakistan into Kimrneridgian belong to Titanosaurs. Titanusaurus does not change the temporal range of the lineage (~Hte Jurassic-Late Meta carpalslmeta tarsals: Four or five Cretaceous) but it expands its spatial distribution brocken .pieces of metacarpals and lor to include Indo-Pakistan. Discoveries of early metatarsals were found in Sun Chako titanosaurids from Africa (Sternfeld, 1911 ; locality .These are rounded to subrounded in Haughton, 1928) and this new discovery from diameter. The thickest specimens diameter Pakistan (Indo-Pakistan subcontinent) and also varies from 3.75 to 5.40 cm. Late Cretaceous advanced Titanosaurids and Saltasurids (Malkani & Anwar, 2000; Malkani Axial Skeleton et al., 2001) have given a new perspective to further attempts to explore and interpret the Ribs: Five brocken pieces of ribs were found in origin and evolution of Titanosauria (Sauropods the Sun Chakko locality. None of them show Dinosaurs). This attempt to update the evolution the tuberculum and capitulum. Three pieces and systematics of the Titanosaurus sauropods is (Fig. 3; MSM-87-K, MSM-92-K, MSM-94-K) based on a comparative analyses of femoral-and belong to the central part of the ribs below the ribs cross sections of early to advanced junction of tuberculum and capitulum. It is Titanosaurids. Fig. 2. Fragments of leg bone like femur/humerus in the upper row. Cross section of femur and fragment of leg bone like femurkmerus in the bottom row (scale is in centimeters).

Fig. 3. Fragments and pieces of ribs in the top row and pieces of rib and proximal fibula in the middle row. Parts of neural spinefdipophysisi zygapophysis and half egghodule concretion/like sexual organ in the bottom row (scale in centimets). 79 I or die ria from the upper Tendaguru beds central part of dorsal ribs cross section vary of Tanzania is usually taken to be the oldest from slightly convex0 to convex (about known member of the Titanosauridae symmetrical) observed in Late Jurassic (Sternfeld, 1911). It is known from several Titanosaurs to plano convex (Asymmetrical) series of very robust limb and foot bones found in Late Cretaceous Titanosaurs and (McIntosh, 1990). The early Cretaceous distal part of dorsal ribs may vary from species from Malawi, " Gigantosaurus" dixeyi plano-convex (symmetrical found in Late (Haughton, 1928) based on caudal, pubis, Jurassic Titanosaurs) to plano-convex scapula and sternal plate, is certainly a (asymmetrical in Late Cretaceous titnosaurid. However, it is not clear whether Titanosaurs). From the above discussion, it it belongs to Tornieria or not (McIntosh, can be found that there was more symmetry 1990). in the ellipticity of femur in Late Jurassic Titanosaurs than found in the Late Cretaceous In Triassic, no definite sauropods Titanosaurs and same is true in the dorsal ribs trackways are known from their time period. cross section. Phylogenetic trend seems like Early and Middle Jurassic is marked by the Brohisaums-Khetranisaud Sulaimanisaurus/ narrow guage tracks, where wide gauge Pakisaurus-Marisaurusl Balochisaurus . trackways are dominant during late Jurassic to late Cretaceous (Wilson et al., 1999). PALEOBIOGEOGRAPHIC IMPLICATION

Primitive sauropods femora show The Kimmeridgian Late Jurassic fauna of Sun eccentricity between the nedio-lateral and Chako and Charroh, Karkh areas, Kirthar antero-posterior femoral but are only slightly Province give the impression of a continuous elliptical (Diplodocus), The femur of relation with the Kimmeridgian fauna of Brachiosaurus has more eccentric profdes , Tanzania and Malawi (Sternfeld, 191 1; and Titanosaurus (Saltasaurus) have the most Haughton, 1928; Weishampel, 1990). As the elliptical sauropod femora. elliptical/eccentric character of cross section of femur and massiveness of leg bones, The tentative evolutionary trends of pneumatic cavities in the anterior ribs and Titanosaurs of Pakistan found in Late Jurassic massiveness in the distal ribs and massiveness (Kimmeridgian) and Late Cretaceous based of metacarpals/metatarsals suggest relation to on morphology of elliptical femoral cross Titanusaurs tentatively named as Brohisaurus section and pieces of dorsal ribs. The general Kirthari. Supposed as the Kimmeridgian evolving trend of femoral cross section started fauna of Pakistan is the same as the one in from circular to oval to suboval to less Tanzania and Malawi, Kimmeridgian elliptical (ancestors) to well elliptical (about Tornieria , the Indo-Pakistan connected with symmetric on antero-posterior axis found in Africa and also belonged to the South Late Jurassic (Kimmeridgian Titanosaurs) to America. However the detailed sampling in well elliptical (asymmetric on antero-posterior the South America and Madagascar are not axis, more thicker on the medial side than the sufficiently known yet to correlate well. lateral side found in the Late Cretaceous Kimmeridgian Titanusaurs found in Pakistan Titanosaur s) as fragmentary and incomplete remains.

The general trend of cross section of During the Late Cretaceous the great dorsal ribs from Late Jurassic Titanosaurs to similarities with the fauna of the South Late- Cretaceous Titanosaurs vary from the America, however, is observed especially in the case of Saltasaurids and Titanusaurids, theropods dinosaurs and Baurusuchids both of them may included into Titanusauria Mesoeucrocodylia. These discoveries of and other archosaurian such as Baurusuchid Jurassic and Cretaceous Arhosaurian reptiles Pabwehshi Pakistanensis (Wilson et al., from Pakistan provide an opportunity for 200 1) and also Vitakridrinda Sulaimi assessing biogeography and phylogeny. Now Abelisaurid Theropod. Pakistan will act as a milestone for the entering of Indo-Pakistan subcontinent into All these Indo-Pakistan subcontinent new hypothesis of Paleobiogeography and genera are distributed in Africa and South Phylogeny. America during Late Jurassic and Early Cretaceous but also distributed in South A land corridor between South America America during Late Cretaceous, Evidence of and Indo-Pakistan is present in the two main close faunistic relation with other continents paleogeographic reconstructions. Smith et . al. and regions has not yet been known may be (1994) reconstruct a corridor between South due to less sampling densitylissue. America and Indo-Pakistan via Antarctica that existed until the Hauterivian (Early At Tendaguru, Tanzania, Titanusaurus Cretaceous) at ca. 130 million years ago. Hay occur (Gigantosaurus =Tornieria) but other et al. (1999) reconstruct the same corridor but Sauropods predominate. Data yet known from propose that it lasted until the campanian the upper Cretaceous of Egypt and the Sahra (Late Cretaceous) at ca. 80 mya. Both depict indicate that the fauna has rather a different' a land connection between Africa and South aspect but it contains a Titanusaurus (Huene America until the Albian-Cenornanian (mid- & Matley, 1933). Cretaceous) at ca. 100 mya, as well as a connection between Indo-Pakistan and Indo-Pakistan initially interlocked with Madagascar that persisted until roughly the the Gondwanan landmasses of Africa, South same time. However, because these two America, Australia, Antarctica and paleocoastline reconstructions indicate Madagascar early in the Mesozoic, drifted different durations for the corridor stretching northward during the Cretaceous to collide between South America and Indo-Pakistan, with Laurasian landmasses during the they predict different landmasses to show Cenozioc . Indo-Pakistan thus appears to have greatest faunal affinity with the Indian experienced a 100 million year period of subcontinent. The paleogeography of Smith et isolation during a 9,000 kilometer migration al. (1994) suggests that Indo-Pakistan's across the equator that can be expected to Cretaceous fauna should display greatest have influenced the character of its native similarity to that of Madagascar, whereas the biota. Fossils from this period of paleogeography of Hay et al. (1999) suggests biogeographic isolation are relatively scarce. greatest similarity to South America and So far, the late cretaceous (Maastrichtian) Antarctica. (Wilson et al., 2001). Lameta formation of India has served as the sole source of information on Cretaceous From the Late Jurassic to Early vertebrates of the Indo-Pakistan subcontinent Cretaceous, the Titanosaurs were dominant in (Wilson et al., 2001). But now Pakistan has a Africa along with the other Sauropods. It variety of large vertebrate such as could not compete the other dominant Titanusaurids and Saltasaurids, sauropodsb S.auropods genera in the Middle and Late dinosaurs, Abelisaurids 1 Tyranosaurids Cretaceous. It may be a less sampling issue. The data likes Kimmeridgian dinosaur in Professor Dr. Philip D. Gingrerich and Dr. Pakistan suggest a direct land communication Jaffery Wilson of the University of Michigan, with the Tanzania and Malawi (African) USA and Professor Ismar S. Carvalho, countries during the kimrneridgian. However, Universidade Federal do Rio de faneiro, dept. in the later time it may isolated from Africa de Geologia/CCMN, Cidade Universitaria- but it remain connected with South America Ilha do Fundao, 21.949-900, Rio de faneiro, through a corridor via Antarctica. Brazil for providing literature on dinosaurs. Paleobiogeographic reconstruction by Smith Thanks are due to Dr. M. Raza Shah, Mr. M. et al. (4994) and Hay et al. (1999) have Faheem Khan of Balochistan University of explained well about the Indo-Pakistan Information Technology and Management subcontinent connection during Late Sciences, Quetta, Mr. Mohammad Ibraheem Cretaceous. But the reconstruction by Hay et Khan of F. G. School, Quetta, Mr. Shireen al. (1999) best accommodates the available Khan, Mr. Ghulam Qadir, Mr. Aqeel Ahmed Late Jurassic and Cretaceous biota of the and Mr. Zahid Hussain of GSP, Quetta for Indo-Pakist an. providing all sorts of help during the accomplishment of this research. CONCLUSIONS REFERENCES The discovery of first Jurassic dinosaur fossils from lowest Sembar Formation, Haughton, S.H., 1928. On some reptilian Kirthar province, Pakistan is of great remains from the dinosaur beds of significance as it has proved that these Nyassaland. Trans. Roy. Soc. S. 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