Fossil biotas from Pakistan with focus on dinosaur distributions and discussion on paleobiogeographic evolution of Indo-Pak Peninsula

M. Sadiq Malkaniand SUN Ge

1. Geological Survey of Pakistan, Quetta, Pakistan; 2. College of Paleontology, Shenyang Normal University, Shenyang 110034, China; 3. Key Lab of Past Life in NE Asia, MRLC, Shenyang 110034, China; 4. Key Lab of Evolution of Past Life & Environment in NE Asia, MOEC, Changchun 130026, China

Fossil biotas from Pakistan with focus on dinosaur distributions and discussion on paleobiogeographic evolution of Indo-Pak Peninsula

M. Sadiq Malkani1and SUN Ge2,3,4

1.GeologicalSurveyofPakistan,Quetta,Pakistan; 2.CollegeofPaleontology,ShenyangNormalUniversity,Shenyang110034,China; 3.KeyLabofPastLifeinNEAsia,MRLC,Shenyang110034,China; 4.KeyLabofEvolutionofPastLife&EnvironmentinNEAsia,MOEC,Changchun130026,China

Recent geological and paleontological exploration in the Indus basin of Pakistan allowed the discoveries of numerous remains of non-marine reptiles (titanosaurian sauropod, abelisaurian and noasaurian theropod dinosaurs), and marine reptiles (crocodiles), flying reptiles (pterosaurs), marine and non-marine mammals, fishes, invertebrates, and plants, especially Pakistan is relatively rich in footprints/trackways in the Mesozoic. These vertebrates of Indo-Pakistan are very significant for paleobiogeographic study due to the present-day connection of this continent with Asia in Northern Hemisphere, whereas during past (Jurassic and pre-Jurassic) it was connected to the Gondwana. The Mesozoic vertebrates show close affinities with Gondwanan landmasses. The Cenozoic vertebrates show Eurasian affinity and migrated from Indo-Pak subcontinent to Eurasia or vice versa via Paleo Indus River systems along Western Indus Suture, after long journey of about 6 000 km the first collision of Indo-Pak subcontinent with Asia occurred at terminal Cretaceous.

fossil biotas, dinosaurs, Pakistan, paleobiogeography, Indo-Pak subcontinet.

1 Introduction

The Indus basin of Pakistan (a Gondwanan fragment) is significant for dinosaur findings as well as Mesozoic terrestrial and marine strata and biotas. This basin is folded and faulted by Cenozoic and end Mesozoic tectonics. The Indus basin shows wide exposures of Mesozoic and Cenozoic marine and terrestrial sequence (Malkani, 2010a) (Fig.1). Due to recent geological and paleontological exploration, Pakistan appeared for the first time on the world dinosaur’s map. Dinosaur research in Pakistan started in 2000 whereas dinosaurs were known from neighbouring India since 1828. All the fossils found so far are housed in the Museum of Geological Survey of Pakistan (GSP) Quetta, Pakistan, except some fossils which are sent to University of Michigan, USA for preparing. The Vitakri region of Pakistan can be considered as a paradise for dinosaurs due to rich fossil bone concentrations. In peace conditions, the application of bulk screen-washing methods of paleontological exploration may reveal the Mesozoic biodiversity of small-sized vertebrates like amphibian (frogs), reptiles (snakes, lizards, turtles, etc), mammals, birds, fishes, etc to make an accurate assessment of the paleobiogeographic affinities of Indo-Pakistan subcontinent

1st Row holotype skull (2 photo) of Gspsaurus pakistani and holotypic skull (4 photo) of Saraikimasoom vitakri of Gspsauridae; 2nd Row holotype (9 photo) of Nicksaurus razashahi of Saltasauridae and holotypic vertebral column (2 photo) of Maojandino alami of Balochisauridae; 3rd Row Trakway (1 photo) of Latest Cretaceous titanosaurs Pashtosaurus zhobi; Trakway (2 photo) of a herd of Middle Jurassic titanosauriforms/early titanosaurs Malakhelisaurus mianwali confronted by a running theropod Samanadrinda surghari; and 5 pieces of titanosaurian sauropod coprolites; 4th Row diverse osteoderms/armor bones of titanosaurs (first in Asia); 5th Row bidiverse limb elements of Pakisauridae and Balochisauridae; 6th Row braincase and atlas-axis of Marisaurus; biconvex first caudal centrum of Balochisaurus; five distal most caudal centrum with trispines (2 photo) making distal ball of different genera; pterosaurs dentaries (2 photo); 7th Row holotypic pair of femora, a braincase, 6 large vertebrae, a whiplash type slender distal caudal centrum of Vitakridrinda and a pes of Vitakrisaurus theropods; 8th Row Mesoeucrocodiles Pabwehshi rostrum articulated with lower jaw; anterior lower jaws of Sulaimanisuchus, snout of Induszalim bala and some postrcranial elements; and mesocrocodiles rib and egg?; 9th Row cranial elements (3 photo) of baluchithere Buzdartherium, jaw (2 photo) of Asifcroco, armour (1 photo) and an egg of eucrocodiles; dentary (3 photo) of cyonids Bolanicyon; 7th Row fish remains from Latest Cretaceous (2 photo) Karkhimachli and Cambrian (a fin) and last 3 photo of Gomphotherium buzdari; 10th Row ammonite impression, nautiloids Pakiwheel, bivalve Pakiring, and a gymnosperm Baradarakht. (Scale each black or white digit is 1 cm)Fig.1 Mesozoic and Cenozoic fossils in Indus basin of Pakistan

A, Distribution of dinosaurs in Pakistan; B, Migration and isolation of Indo-Pak plate during Cretaceous; C, Paleo-Indus River systems flow from north to south and northwest to southeast; D, Paleo-Vitakri River systems generally flows from East to west; E, Distribution of dinosaurs in Indo-Pak subcontinent.Fig.2 Distributions of dinosaurs in Pakistan and their related geological background in Indo-Pak plate during Jurassic-Cretaceous

(Malkani & Sahni, 2015). These diverse fossils and their sites need to be protected by creating national and global Geoparks for tourism which will be an innovation for the development of provinces of Pakistan and also useful for world science. All these materials have great potential to resolve lower and higher level phylogeny, paleobiogeographic link, tectonics evolution and geodynamics of Indo-Pakistan subcontinent which was isolated from Madagascar and other Gondwanan landmasses, migrated (more than 6 000 km) from south to north and collided with Asia (Fig.2). The data presented here correspond to about 3000 fossilized bony remains of saurischian dinosaurs, mesoeucrocodiles, pterosaurs and some footprints and trackways of dinosaurs from Mesozoic; and some new faunas and floras from Cenozoic of Pakistan that have been discovered and collected by the first author and his colleagues.

2 Fossil biotas in Pakistan

2.1 Dinosaurs

(1) Titanosaurian sauropod dinosaurs from Jurassic-Cretaceous

BrohisauruskirthariMalkani 2003, a titanosauriform or early titanosaurian sauropod found in the Late Jurassic lowest part of Sembar Formation from Sun Chaku locality of Karkh area of Kirthar Range and it is based on some poorly recognized postcranial elements (Malkani, 2003a). Only one piece of bone from Lakha Kach Charo Zidi area is referred.

KhetranisaurusbarkhaniMalkani 2004, a pakisaurid titanosaurian sauropod (slightly broad to sub squarish midcaudals; based on one holotypic vertebra collected from mid Kinwa (northwest of stream) and one attributed caudal from Sangiali (Malkani, 2006), and three attributed caudal from Bor, Mari Bohri and Grut Gambrak localities (Malkani, 2009).

SulaimanisaurusgingerichiMalkani 2004, a pakisaurid titanosaurian sauropod based on seven fragmentary but associated caudal vertebrae (long and squarish mid caudals) collected from Kinwa locality (Malkani, 2006), and referred vertebrae belong to Shalghara and Mari Bohri localities (Malkani 2008a, 2009, 2010c), one distal femur, and 100 axial and appendicular associated fossils from Sangiali locality.

PakisaurusbalochistaniMalkani 2004, a pakisaurid titanosaurian sauropod based on four associated tall caudal centra (Malkani, 2006) associated with many surface finds of postcranial skeletons like, vertebrae, ribs, sternal, scapulae, humerai, radius, ulnae, ilia, femora, tibia, fibula, foot bones, osteoderms, pieces of coprolite, etc. All fossils are collected from the latest Cretaceous Vitakri Formation in Kinwa, Bor, Top Kinwa, Shalghara, Sangiali, Alam Kali Kakor (north), Zubra, Darwaza, and other localities in Vitakri area, Barkhan, Balochistan (Malkani 2008a, 2010a,c, 2014a, 2015a,d).

MarisaurusjeffiMalkani 2004, a balochisaurid titanosaurian sauropod, based on six heavy and slightly tall to squarish midcaudals (Malkani 2006). It is characterized by a relatively short and thick tail with trispinous distalmost tail special. It is relatively stocky and enormous. Its first caudal centrum is biconvex. It is attributed by cranial and postcranial materials (Malkani, 2008a,b, 2010c, 2014a, 2015a,d; Wilsonetal., 2005).

BalochisaurusmalkaniMalkani 2004, a balochisaurid titanosaurian sauropod and based on seven associated heavy and broad to squarish caudal vertebrae (Malkani, 2006). It is a most advanced and large-sized dinosaur characterized by a relatively more stocky and large body with stocky limbs. Its first caudal centrum is biconvex. It has a short and stocky tail with distal trispinous ball distalmost end of tail.

GspsauruspakistaniMalkani 2014, a large-sized gspsaurid titanosaurian sauropod found from red muds of Vitakri Formation of Alam Kali Kakor locality of Vitakri area and based on a mostly complete skull (Malkani, 2014a, 2015a,d). It represents pneumatic skull, V shape palatal hook and broad V shape palatal shelf, long, slender, slightly recurved and sub circular to oval teeth, thick dentary and jaw rami. The anterior portion of upper jaw is broadly arched forming U shape and lower jaw forming V shape. Tooth slenderness index varies from 4--5.

SaraikimasoomvitakriMalkani 2014, a young gspsaurid titanosaurian sauropod based on a mostly complete skull (Malkani, 2014a, 2015a,d). Its rostrum represents 400inclinations from the horizontal. It represents relatively stocky, straight and subcircular teeth, slender dentary and jaw ramus and pneumatic skull bones. The anterior portions of upper and lower jaws are broadly arched forming U shape. Tooth slenderness index varies from 3--5.

NicksaurusrazashahiMalkani 2015, it is based on axial and stocky limb elements of young adult of this taxon from Kinwa north locality of Vitakri area include a pair of femora (left full femur, right partial femur), a pair of stocky distal tibiae, partial humerus parts; proximal radius, five teeth in jaw ramus, teeth, skull fragments, chevron, cervical, dorsal and caudal centra (Malkani, 2014a, 2015a,d).

MaojandinoalamiMalkani 2015, a large bodied balochisaurid titanosaurian sauropod (Malkani 2015a) with a thick, broad and long neck, short tail, heavy and stocky body and it is based on associated holotypic materials including about 6 cervical, 4 dorsal, and 10 caudal vertebrae along with partial left femur, partial left and right tibiae, and partial radius, a pair of partial distal scapulae, partial sternal plate or ilia, some neural arch and laminae covered partially by yellow brown muds, etc. The holotypic axial and limb elements collected from Alam Kali Kakor locality of Vitakri area.

Trispinous distal tail ball of titanosaurs

The distalmost caudal centrum of some Pakistani titanosaurs provide a new look like a trispinous distal tail ball (Malkani, 2008a, 2012c, 2015a,d) used for multipurposes like good defending tool against its foe, balancing body as third support during foraging from tall tree, and mating. Tail of titanosaurs from Pakistan is unique in the world. This distal most caudal centrum and their spines are compact and not cancellous. So far this discovery shows an endemic shape of Pakisauridae and Balochisauridae.

Osteoderms and dermal plates of dinosaurs

Osteoderms of titanosaurs from Pakistan are reported first time in Asia (Malkani 2003b). The six types of titanosaurian osteoderms are reported so far from Pakistan (Malkani, 2003b, 2010c, 2015d). The first type is a simple plate (compact) without mosaic in nature, the second is a mosaic plate (cancellous), the third type is a large ellipsoid (without median cut) with cone (cancellous), the fourth is a large ellipsoid (with median cut) with cone (cancellous), the fifth is an ungual with cone (compact) and the sixth type is a jaw with teeth type (cancellous).

Coprolites of titanosaurian sauropod dinosaurs

Coprolites of dinosaurs are found from Pakistan with some wheel type rounded and spongy/vesicular pieces with 5--7cm in diameter are found along with bones of titanosaurian sauropods from the red mud horizons of the Latest Cretaceous Vitakri Formation in the Vitakri dome area (Malkani, 2015d).

(2) Theropod dinosaurs from the latest Cretaceous Vitakri Formation

VitakridrindasulaimaniMalkani 2004, a large sized carnivorous abelisaurian theropod dinosaur and based on a pair of left and right proximal femora and basioccipital condyle along with partial braincase (Malkani 2006a, 2014b). The femur has a cavity hollow with thick walls as present in most theropods, while the vertebrae may belongs to more than multiple large theropods present in the Vitakri Formation. Its massive and thick braincase parietal/ dorsal cover matches withCarnotaurussastreiandIndosaurusmatleyi, its basipterygoid process is high angle while inIndosuchusraptoriusis low angle (M. Mortimer, personal communication) The foramen magnum is covered by matrix in the posterior view (Malkani, 2014b, 2015d).

VitakrisaurussaraikiMalkani 2010, a small sized carnivorous Vitakrisauridae noasaurian theropod and based on isolated pes collected from the terrestrial overbank red muds of latest Cretaceous Vitakri Formation in Bor Kali Kakor, Vitakri area (Malkani, 2010b). The foot is tridactyl. The phalanges of right hind limb are robust. There are three preserved digits. According to M. Mortimer (personal communication, 2012) the taxon is more comparable to noasaurids than abelisaurids based on its size. The large ratio between phalanges II-2 and II-1 and proximally narrow metatarsal II (in dorsal view) also suggest noasaurids affinities while phalanx II-2 is longer thanVelocisaurus(compared to phalanx II-1), this varies within other species, and further comparison is not possible pending better description of theVitakrisaurusand other noasaurid pes (M. Mortimer, personal communication, 2012).

(3) Trackways of sauropod and theropod dinosaurs

For the first time in Indo-Pakistan a footprint of sauropod dinosaur was discovered from the Upper Cretaceous of India. Then trackways of titanosauriform or early titanosaurian sauropodMalakhelisaurusmianwaliMalkani 2008 herd and a running theropodSamanadrindasurghariMalkani, 2008 were discovered from the Middle Jurassic Samanasuk Limestone of Kohat and Potwar basin (Malkani, 2007, 2008a). An isolated footprint of sauropod was also found 1 km southward from theMalakhelisaurussite in the same formation. Further 400 m southward from the above mentionedSamanadrindasite, tracks of a couple of small bodied noasaurian theropodsHimalayadrindapotwariMalkani 2015 were found from the Middle Jurassic Samanasuk limestone of Kohat and Potwar basin (Baroch Nala, Malakhel area, Mianwali District, Punjab, Pakistan). The new footprints and trackways of titanosaurian sauropodPashtosauruszhobiMalkani 2014 are found on the Sandstone beds of Vitakri Formation of Qila Saifullah District, Balochistan. There is s danger that these trackways may be destroyed during future construction materials required for road construction. The large oval manus footprints of titanosaurs from the Upper Cretaceous (Malkani, 2014a, 2015c) reveal that the titanosaur’s manus were without claws. Further the large footprints about 1m of pes of titanosauriforms from Middle Jurassic (Malkani, 2008a) and titanosaurs from the Upper Cretaceous (Malkani, 2014a, 2015b,d) reveals that the titanosaurs pes were tripodials and other two digits are highly reduced or lost.

2.2 Pterosaurs

SaraikisaurusminhuiMalkani 2013, a pterosaur found from the latest Cretaceous Vitakri Formation of Top Kinwa, Vitakri area (Malkani, 2013, 2014b). The preserved dentary ramus shows carnivorous type elongated skull with eight teeth. This ramus shows internal pneumatic texture and structure. The teeth are oval to suboval, some overlapped.

2.3 Crocodiles

PabwehshipakistanensisWilsonetal. 2001 is based on rostrum and articulated partial dentary.

SulaimanisuchusskinwaiMalkani 2010 is based on anterior dentary and splenial. The first and third tooth are small in diameter while third is relatively more but the fourth is maximum in dia, the fourth dentary teeth is large and shows a marked heterodonty in size and is mostly compressed (Malkani 2014b, 2015b).

InduszalimbalaMalkani 2014, a terrestrial mesoeucrocodile Induszalimidae taxon, based on rostrum found from mid Alam locality (Malkani, 2014b, 2015d). InInduszalim, the snout depth is equal to width(inPabwehshithe depth is 3/4 of width), the maxillary ramus is relatively far away/down to palatal contact (inPabwehshithe maxillary ramus is just close to palatal contact), the maxillary ramus on the lateral side has pits with anteroposterior lineation (Pabwehshihas no lineation), and there is marked boundary/contact between palatal shelves and maxillary wall (inPabwehshino boundary/contact shown). Moreover, inInduszalimthe large teeth dia is relatively larger thanPabwehshiin tooth diameter, andInduszalimmatches with thePabwehshion saggital torus on palatal shelves ofPabwehshi.

KhuzdarcrocozahriMalkani 2015d is based on partial rib found from the maroon shale and marl alternated beds of Early to Middle Cretaceous Goru Formation near Khuzdar town. Furthermore, one possible egg fragment may belong toKhuzdarcrocozahrior any other genera of mesocrocodile/ mesoeucrocodile, which is found from the Cretaceous of Karkh, Khuzdar District, Balochistan. The egg is amorphous in inner portion, preserved as 4 cm long by 3.8--4.2 cm in diameters.

2.4 Fish

The possible bony fishes, the Teleostei or holostei fish or ichthyosaurKarkhimachlisangialiare found fragmentary on the Early Paleocene part of Sangiali Group of Karkh area of Khuzdar District but its nearby higher areas consists of Late Cretaceous Mughalkot and Pab formations. Its age is being assumed as Late Cretaceous. Its preserved portion mostly belongs to body cross section having herring bone type structure. It is small sized fish/ichthyosaur. Further some cross sections are also referred to it. A body cross section of marine fish found in the Jurassic Chiltan Limestone of Kharzan of Mula-Zahri area, Kirthar Range Malkani 2014c. Further a fin/partial cross section of Cambrian fish found as fragmentary in the dolomitic limestone of Azad Kashmir (Malkani, 2015d).

2.5 Eucrocodiles

A eucrocodileAsifcrocoretraiMalkani 2015b is found in the ferruginous conglomeratic bed of Oligocene Chitarwata Formation in Gulki area (Taunsa-Gulki section) of Dera Ghazi Khan District (eastern Sulaiman foldbelt). It is fresh water crocodiles living in the Paleo Indus River. Its centrum is opisthocoelous type having ball on anterior and concave cavity/coel in the posterior. Its ulna have dorsal groove to set humerus. Further the three eggs are collected, one egg may belongs to crocodiles while other egg may belongs to birds/snake/lizard. This egg like material may be the osteoderms/coprolite of baluchithere because these are found in the association with theBuzdartheriumgulkiraoMalkani, 2015d.

2.6 Mammals

PakitheriumshagalaiMalkanietal. 2013-a baluchithere reported first time in Balochistan basin.BuzdartheriumgulkiraoMalkani 2014c from Taunsa area of Dera Ghazi Khan District (east and northern Sulaiman) is the first time discovery of baluchithere in Punjab Province and third locality of baluchitheres in Pakistan after Dera Bugti and Shagala. These hornless rhinos Baluchithere were widespread during the Eocene to Oligocene in Asia (Pakistan, China, Mongolia, Kazakhstan) and southeastern Europe (Yugoslavia, Bulgaria, Romania, Turkey Georgia). The bones/pieces of bones belong to cranial, vertebral and limb elements. The tusk like incisor, large size of the bones, large size of ungual/toe, their close occurrences and no duplication tells their association with one individual of baluchithere (Malkani 2014c, 2015d).

SulaimanitheriumdhanotriMalkani et al. 2013 is third genus and species of Basilosauridae (king of basal whale) in Asia. It is important due to present discovery of almost complete vertebral column (Malkanietal., 2013). The walking whale from Pakistan tells the evolution from land to sea (Gingerichetal., 2001).

GomphotheriumbuzdariMalkani 2014 a big proboscidean remains collected from Miocene Litra Formation of Vihowa group in the Zinda Pir anticline area located in the easternmost extremity of Sulaiman fold and thrust belt (Malkani, 2014c, 2015d).

BolanicyonshahaniMalkani 2014 (Quettacyonidae) is found at the coal mining at depth of about 500 m in the Early Eocene Toi Formation of southwestern Mach (Gishtari) area. Its 1 incisor, 1 canine, 4 premolar and 3 molar teeth are preserved. This mammal is among the oldest in Pakistan and it may be an old horse (Malkani, 2015d). Pakistan has a unique blend of Tertiary mammals of Asian affinity like rhinocerotoids and antharacotherids with few proboscideans, carnivores, chalicotheres, deinotheres, bovids, suids, Creodonta, ruminantia, amynodontiae, and crocodilians (Eames, 1970; Raza & Meyer, 1984).

2.7 Invertebrates

The invertebrates including Protozoa, Foraminifera, Coelenterata, Stromatoporoidea, Scaphopoda, Bryozoa, Brachiopoda, Fusilinids, Echinodermata, Conodonts, Mollusca, Cephalopoda, Gastropoda, Pelecypoda/Bivalves and Arthropoda, have been reported from Pakistan.Pakiringkharzani(Malkani, 2014c) (bivalves) is found from the Cretaceous-Paleogene (K-Pg) boundary laterite/thin rust on the last bed of Pab sandstone in the Kharzan area of Khuzdar district. It is sub ring type and rough surface ornamented bivalves with rope like shape.Mulastarzahri, a star fish, two pectin type other bivalves, 3 gastropods and 1 coral like fossils are found from Shaheed Ghat Formation of Kharzan area, Mula-Zahri range.Pakiwheelvitakri, the stocky type nautiloids is found just after the K- Pg boundary in Sangiali Formation close to east of Vitakri town (Malkani, 2014c), andPakiwheelkarkhi, the slender type nautiloids, is found in the green mudstone, may be of volcanic origin, of the Early Paleocene Sangiali Formation, 5 km east of Karkh town (Malkani, 2015d).

2.8 Cretaceous plants

The fossil wood, gymnosperm and angiosperm pollen, and fern spores have been reported from the Cretaceous of Pakistan. A conifer wood log ofBaradarakhtgoeswangaiMalkani, with 20 cm in diameter, is found from the Latest Cretaceous (Maastrichtian) Pab Formation in Goeswanga Pass, Barkhan District, Balochistan (Malkani, 2014c). More fossil plants may be found further from the Kingri coal of Vitakri Formation (Malkani, 2014c).

3 Dinosaurian distributions in Indo-Pak Peninsula

The dinosaur evidences from Pakistan are found in Indus basin which is further subdivided into Upper Indus (Kohat and Potwar), Middle Indus (Sulaiman) and Lower Indus (Kirthar) basins. In the Lower Indus (Kirthar) basin, a few and poorly recognized body fossils of /titanosauriforms/early titanosaurian sauropod were found from the Late Jurassic Sembar Formation of Karkh area, Khuzdar District, and a footprint of a sauropod in the Middle Jurassic chiltan Limestone. In the Upper Indus (Kohat and Potwar) basin, trackways of a herd of more wide gauge titanosaurian sauropods confronted by a running narrow gauge theropod and a track of a couple of small theropods and a single pes print of titanosauriform are reported from the Middle Jurassic Samanasuk Limestone of Baroch ichno type, Malakhel area, Mianwali District, Punjab. The Middle Indus (Sulaiman) basin yields the latest Cretaceous body fossils of titanosaurian sauropods, aabelisaurian and noasaurian theropods, mesoeucroccodiles and pterosaurs, and titanosaurian sauropods trackways (Malkani, 2015d). In India dinosaurs are found from the states of Rajasthan and Gujarat (Western sector), Madhya Pradesh and Maharashtra (Central sector) and Andhra Pradesh, Tamil Nadu and Karnataka (Southern sector) (Sahni, 2001). India is relatively rich in eggs and nests, Triassic and Jurassic fossils while Pakistan is relatively rich in footprints/trackways and Latest Cretaceous fossils (Malkani and Sahni, 2015).

4 Paleobiogeographic evolution of Indo-Pak Peninsula

Indo-Pak is a peninsula but in the past it was a part of Gondwanalands, and during northward journey it was an island (Fig.2). During the Late Triassic (ca. 220 Ma) the lands united as Pangea. The breakup of Pangaea started in the Middle Jurassic (170 Ma) while the breakup of Gondwana started in the Late Jurassic (160 Ma). Indo-Pakistan separated from Madagascar during the Late Jurassic and started northward journey during the Early Cretaceous (135 Ma). Most of Cretaceous passes as journey and island. It migrated rapidly covering more than 6 000 km in about 68--70 million years ago. Its northwestern corner collided first time with Afghan block of Tethys and Asia at the latest Cretaceous about 67 Ma ago. This corner acts as a pivot point for counterclockwise rotation. This first collision (67 Ma) created the land bridge (between Indo-Pakistan Shield and Asia) from Afghanistan/Hindu Kush-Kurram-Waziristan-Zhob (Western Indus Suture and adjoining Afghan land)-Ziarat-Fort Munro (D. G. Khan) areas, allowed the migration of the latest Cretaceous fauna from Asia to Indo-Pakistan and vice versa. During the Paleocene the sea transgressed on this land bridge and other vicinity areas and deposited the marine Sangiali Group in the Sulaiman basin and Nisai group in the Balochistan basin. During the late Paleocene the sea regressed from the western Sulaiman basin and northern Balochistan basin due to further uplift and collision by continued movements of subcontinent. Consequently the Western Indus Suture, close to the Zhob-Waziristan-Kurram and its adjoining western areas of Afghan block uplifted more. This uplift provided the lands for the migration of the Eocene vertebrates like baluchithere-the largest rhinoceros from Asia to Indo-Pakistan subcontinent or vice versa. This rising ended the Paleo Vitakri River systems of Sulaiman foldbelt flowing from east to west (Indo-Pakistan shield toward Neotethys) and started the birth of Paleo Indus River systems flowing from northwest to southeast and north to south. It deposited the deltaic and terrestrial molasse Late Paleocene-Early Eocene Chamalang (Ghazij) Group in the Sulaiman basin, Shagala Group of Balochistan (fore arc or arc-trench gap, arc, back arc) basin. This molasse shows the beginning of Neotethys closure from NW of Indo-Pakistan subcontinent. The middle Eocene is represented by a major transgression of sea in the westward and also eastward. As a result the transition period of this transgression is represented by the deposition of upper Ghazij Group (Drug limestone and Baska evaporitic gypsum, and shale show rhythmic off and on sea water) and major transgression is represented by the deposition of Kahan Group (Habib Rahi limestone, Domanda shale, Pirkoh limestone and Drazinda shale) in the Sulaiman basin of Pakistan while the northern Balochistan/Kakar Khorasan basin was under erosion. At the latest Eocene (40--35 Ma) the northern part of Indo-Pakistan collided with Karakoram and Tibetans parts of Asia. This collision resulted in the uplift, folding and faulting (mainly south verging thrusts) in the northwestern Foreland (Hindukush-Karakoram belt and Kohistan-Ladakh magmatic arc) and its adjoining northern part of Hinterland now called Himalaya. Consequently Tethys Sea was permanently closed by the end Eocene tectonic episode which is responsible for the birth of Himalaya. This collision started the terrestrial/continental fluvial facies and Neotethys permanently closed from Hindu Kush-Karakoram belt, Kohistan magmatic arc, Khyber-Hazara (uppermost Indus), Kohat and Potwar (northern/upper Indus) and Sulaiman (Middle Indus) basins, while the Kirthar (southern/lower Indus) basin was remained undersea as the Oligocene marine Nari and Gaj formations were deposited in Kirthar basin. This episode/uplift increases the gradient of Indus River generally flowing from north to south and also created the birth of Ganges River systems generally flowing from west to east and northwest to southeast. This uplift is responsible for the molase deposition of Siwalik, Potwar and Vihowa groups in Indo-Pakistan Subcontinent. Consequently the Neotethys closed from the area presently called Himalaya. The main geoevents occurred at the Pliocene-Pleistocene boundary time resulted for the further retreat of sea from the Kirthar (lower/southern Indus) basin as the terrestrial Vihowa/Manchar Group deposited. So far the last major episode occurred at the end Pleistocene, resulted further retreat of sea in the south and folding and faulting (Malkani, 2011; Malkani & Mahmood, 2016). The northward movements of Indo-Pakistan plate are continuing so far. This tectonic orogeny is responsible for creating highest peaks and present morphology. The Neotethys remained in the east, south and mostly in the west of Indo-Pakistan subcontinent but now named as Indian Ocean. Due to the closure of the Neotethys from north, northwest and some part of west of Indo-Pakistan subcontinent, shaped it as Peninsula. Due to contact and collision of Indo-Pakistan subcontinent with the southern part of Asia it is being called as South Asia. Mesozoic vertebrates show close affinities with Gondwanan landmasses. The Cenozoic vertebrates show Eurasian affinity and migrated from Indo-Pak subcontinent to Eurasia or vice versa via Paleo Indus River systems along the Western Indus Suture, after the first collision of the Indo-Pak subcontinent with Asia occurred at terminal Cretaceous, however the baluchithere-the largest land mammals show Indo-Pak origin. The present paleobiogeographic study would also be beneficial for the related study in China, particularly for the study in the Tibet and Xinjiang regions (Sun, 1987, 1993; Sunetal., 2014; Martinetal., 2010).

Eames F E. 1970. Some thoughts on the Neogene/Paleogene boundary.Paleogeogr.Paleoclim.Paleoecol.,8: 37-48.

Gingerich P D, Haq M U, Zalmout I,etal. 2001. Origin of whales from early artiodactyls: Hands and feet of Eocene Protocetidae from Pakistan;Science, 295: 2239-2242.

Malkani M S. 2003a. First Jurassic dinosaur fossils found from Kirthar range, Khuzdar District, Balochistan, Pakistan.GeologicalBulletinUniversityofPeshawar, 36: 73-83.

Malkani M S. 2003b. Pakistani Titanosauria are armoured dinosaurs?.GeologicalBulletinUniversityofPeshawar, 36: 85-91.

Malkani M S. 2004. Saurischian dinosaurs from Late Cretaceous of Pakistan//Hussain S S, Akbar H D. (Eds.) Fifth Pakistan Geological Congress, National Geol. Society Pakistan, Pakistan Museum of Natural History (Pak Sci Foundation), April 14-15, Islamabad, Pakistan, 71-73.

Malkani M S. 2006. Biodiversity of saurischian dinosaurs from the latest Cretaceous Park of Pakistan.JournalofAppliedandEmergingSciences, 1(3): 108-140.

Malkani M S. 2007. Trackways evidence of sauropod dinosaurs confronted by a theropod found from Mid Jur. Samana Suk Limestone of Pakistan.SindhUniversityResearchJournal:ScienceSeries, 39(1): 1-14.

Malkani M S. 2008a.Marisaurus(Balochisauridae, Titanosauria) remains from the latest Cretaceous of Pakistan.SindhUniv.Res.Journ.:Sci.Series, 40(2): 55-78.

Malkani M S. 2008b. First articulated Atlas-axis complex of Titanosauria (Sauropoda, Dinosauria) uncovered from the latest Cretaceous Vitakri member (Dinosaur beds) of upper Pab Formation, Kinwa locality of Sulaiman Basin, Pakistan.SindhUniversityResearchJournal:ScienceSeries, 40(1): 55-70.

Malkani M S. 2009. NewBalochisaurus(Balochisauridae, Titanosauria, Sauropoda) andVitakridrinda(Theropoda) remains from Pakistan.SindhUniversityResearchJournal:ScienceSeries, 41(2): 65-92.

Malkani M S. 2010a. NewPakisaurus(Pakisauridae, Titanosauria, Sauropoda) remains and Cretaceous- Tertiary (K-T) boundary from Pakistan.SindhUniversityResearchJournal:ScienceSeries, 42(1): 39-64.

Malkani M S. 2010b. Stratigraphy and Mineral potential of Sulaiman (Mid. Indus) basin, Pakistan.SindhUniversityResearchJournal:ScienceSeries, 42(2): 39-66.

Malkani M S. 2010c. Osteoderms of Pakisauridae and Balochisauridae (Titanosauria, Sauropoda, Dinosauria) in Pakistan.JournalofEarthScience, 21(Spec. 3): 198-203.

Malkani M S. 2013. New pterosaur from the latest Cretaceous terrestrial strata of Pakistan//Abstract Book of 9thInternational Symposium on the Cretaceous System, September 1-5, Metu Congress Center, Middle East Technical University, Ankara, Turkey, 62.

Malkani M S. 2014a. Titanosaurian sauropod dinosaurs from the latest Cretaceous of Pakistan//Abstract Volume of 2ndsymposium of International Geoscience Program 608 (IGCP 608) “Cretaceous Ecosystem of Asia and Pacific”, Tokyo, Japan, 108-111.

Malkani M S. 2014b. Theropod dinosaurs and mesoeucrocodiles from the Terminal Cretaceous of Pakistan//Abstract Volume of 2ndsymposium of International Geoscience Program 608 (IGCP 608) “Cret. Ecosys. of Asia & Pacific”, Tokyo, Japan, 169-172.

Malkani M S. 2014c. Records of fauna and flora from Pakistan; Evolution of Indo-Pakistan Peninsula//Abstract Volume of 2ndsymposium of International Geoscience Program 608 (IGCP 608) “Cretaceous Ecosys. Asia and Pacific” Tokyo, Japan, 165-168.

Malkani M S. 2015a. Titanosaurian sauropod dinosaurs from Pakistan//Zhang Y. (ed.) Abstract volume, 12thSymposium on “Mesozoic Terrestrial Ecosystems (MTE 12), and 3rdSymposium of International Geoscience Program (IGCP 608) “Cretaceous Ecosystem of Asia and Pacific” Shenyang, China, 93-98.

Malkani M.S. 2015b. Terrestrial mesoeucrocodiles from the Cretaceous of Pakistan//Zhang Y. (ed.) Abstract volume, 12thSymposium on Mesozoic Terrestrial Ecosystems (MTE 12), and 3rdSymposium of International Geoscience Program (IGCP 608) “Cretaceous Ecosystem of Asia and Pacific” Shenyang, China, 242-246.

Malkani M S. 2015c. Footprints and trackways of dinosaurs from Indo-Pakistan Subcontinent-Recent Advances in discoveries from Pakistan//Zhang Y. (ed.) Abstract volume, 12thSymposium on Mesozoic Terrestrial Ecosystems (MTE 12), and 3rdSymposium of International Geoscience Program (IGCP 608) “Cretaceous Ecosystem of Asia and Pacific” Shenyang, China, 186-191.

Malkani M S. 2015d. Dinosaurs, mesoeucrocodiles, pterosaurs, new fauna and flora from Pakistan. Geological Survey of Pakistan, Information Release 823: i-iv,1-32.

Malkani M S, Dhanotr M S I, Latif A,etal. 2013. New remains of Basilosauridae-the giant basal whale, and baluchithere-the giant rhinoceros discovered from Balochistan Province (Pakistan).SindhUniversityResearchJournal:ScienceSeries, 45(A-1):177-188.

Malkani M S, Mahmood Z. 2016. Revised stratigraphy of Pakistan. Geol. Surv. Pak, Record 127: i-iv,1-87.

Malkani M S, Sahni A. 2015. A unified perspective of Terrestrial Jurassic and Cretaceous vertebrates from the Indian Subcontinent: Challenges and Prospects//abstract volume of International conference on “Current perspectives and emerging issues in Gondwana evolution”. Lucknow, India, 11.

Martin T, Sun G, Mosbrugger V. 2010. Triassic-Jurassic biodiversity, ecosystem, and climate in the Junggar Basin, Xinjiang, Northwest China.Palaeobio.Palaeoenv., 90: 171-173

Raza S M, Meyer G E. 1984. Early Miocene geology and paleontology of the Bugti hills, Pakistan//Shah S M I, Pilbeam D. (Eds.) Cont. Geology of Pakistan. Geol. Surv. Pakistan, Mem., 11: 43-63.

Sahni A. 2001. Dinosaurs of India. Delhi: National book Trust, 1-110.

Sun G. 1987. On Late Triassic geofloras in China and principles for palaeophytogeographic regionalizations.ActaGeologicaSinica, 61: 1-9.

Sun G. 1993. The Late Triassic flora from the Tianqiaoling of Wangqing, China. Changchun: Jilin Sci. Techn. Press, 1-157.

Sun G, Akhmetiev M, Markevich V,etal. 2014. The Cretaceour-Paleogene (K-Pg) boundary from Jiayin of Heilongjiang, China and the extinction of dinosaurs. Shanghai: Shanghai Sci. Techn. Educ. Press, 1-194.

Wilson J A, Malkani M S, Gingerich P D. 2001. New Crocodyliform (Reptilia, Mesoeucrocodylia) form the upper Cretaceous Pab Formation of Vitakri, Balochistan (Pakistan),ContributionsformtheMuseumofPaleontology,TheUniversityofMichigan, 30(12): 321-336.

Wilson J A, Malkani M S, Gingerich P D. 2005. A sauropod braincase from the Pab Formation (Upper Cretaceous, Maastrichtian) of Balochistan, Pakistan.GondwanaGeol.Mag., Spec.Vol. (8):101-109.

10.3969/j.issn.1673-9736.2016.04.04

1673-9736(2016)04-0230-11

Received 30 September 2016, accepted 28 October 2016