PINSA 65. A. No. 3, May 1999, pp. 397-425 © Printed in India. THE BREAKUP OF GONDWANA AND THE EVOLUTION AND BIOGEOGRAPHY OF THE INDIAN PLATE SANKAR CHATTERJEE AND CHRISTOPHER R SCOTESE* ‘Muscum of Texas Tech University, Box 43191, Lubbock, TX 79409-3191 (USA) *Department of Geology. University af Texas. Box 19049, Arlington. TX 76019-0049 (USA) (Received and Accepted 14 May 1999) ‘The palacopositions of India afier its breakup from Gondwana and its subsequent northward journey during the Mesozoic and Early Palacogene pose many plate tectonics and palaeobiogeographic riddles. Most reconstructions show peninsular India separating from Gondwana, and remaining an island continent for more than 100 million years until it collided with Asia. However. the lack of endemism among Indian Cretaceous terrestrial vertebrates is, clearly inconsistent with the island continent hypothesis. A new model for the tectonic evolution of the Indian plate Irom its Pangean ongin to the present day 18 proposed that 1s well constrained by geological and geophysical evidence. In this model, a previously unrecognized land area, called Greater Somalia, occupied the position between eastém Arabia and northwestern India, During the Late Jurassic, India rifted away from Greater Somalia. Throughout most of the Cretaceous India was separated from the rest of Gondwana but in the latest Cretaceous it reestablished contact with Africa through Greater Somalia. India maintained contact until the Eocene Period when it colfided with Asia. Based on this plate tectonic model, 14 new paleogeographic maps are presented showing the evolution of the Indian plate from its Pangean origin to its Gal union with Asia. An area cladogram identifies the 11 nodes of hierarchial tectonic evolution of the Indian plate during the last 250 million years. ‘A major question of Indian palaeobiogeography is how the terrestrial vertebrates such as dinosaurs responded to ‘ragmentation of India from Gondwana. The similarities and citterences between Indian vertebrates with those ot other continental fragments provide an indepesdent check for a number of key events during its tectonic evolution. ‘Using the plate tectonic reconstructions as a guide, the distributions of ginosaurs and other vertebrates during the Mesozoic and Early Cenozoic are discussed. A close correspondence between the continental position of India and the distribution of vertebrates is generally observed. However, the Indian vertebrate fauna seems more closely related to Europe at the famitial level to that of Africa during the Mesozoic. This disjunct endemism of fossil distribution pattern may reflect poor sampling from the intervening Africa. Whenever Indian vertebrate records are missing from certain geologic periods, their compositions are inferred from the contemporary animals in adajacent landmasses, ‘The Late Triassic and Jurassic vertebrates of India are cosmopolitan in the Pangean world indicating various trans- continental migration routes. Four such dispersal routes are identified, northern, central, southern and westem. During the mid-Cretaceous, India became isolated and these dispersal routes were closed. During the Late Cretaceous (approximately 70 Ma), the northem route across the Greater Somalia was reopened allowing immigration of dinosaurs and other vertebrates from Africa and Europe. This northern dispersal roure explains for the first time why the Indian Maastrichtian vertebrates do not shaw any evidence of endemism. About 65 million years agy, the Shiva impact evcut at he Indu-Seychelles buundary and the Chivaulul impact in Mexivy tiggered 3 biotic catastrophe and led to the demise of dinosaurs and other organisms. Mammals rebounded from this catastrophe and became the dominant land vertebrates during the carly Tertiary. Fossil evidence suggests that India ‘made the initial contact with Asta during the Eocene and opened a new northeastern corridor for faunal interchange. Key Words: India; Plate Tectonics; Pangea; Gondwana; Dinosaurs; KT Extinction; Biogeography; Shiva Crater; Plate Tectonic Reconstructions Introduction tum affects climate, environment, food supply, speciation, and the diversity of life. Dinosaurs The evolution of life is intricately linked to the (meaning nonavian dinosaurs throughout the text) evolution of lithosphere. Plate tectonic movements provide an interesting case study to understand create mountains and oceans, break and suture how evolution is affected by shifting continents continental fragments, alter geography, which in and changing environments. Dinosaurs first308 appeared in the Late Triassic when most continents were joined together to form Pangea. Their radiation and diversification in the Jurassic and Cretaceous periods were greatly controlled by the fragmentation of Pangea and opening and closing of new biogeographic corridors across the separated landmasses. The coincidence of dinosaur diversification and continental fragmentation provides a new arena for testing various biogeographic models such as vicariance, dispersal, speciation, endemism, homogenization, and phylogeny. ‘he distribution of dinosaurs trom Alaska to Antarctica suggests that climate may not be significant deterrents to their migration. On the other hand, dinosaurs were exclusively terrestrial animals and deep, wide oceanic tracts must have been an effective barriers to migration. When Pangea broke up, the cosmopolitan dinosaur populations were isolated from each other, leading to genetic isolation and increasing endemism. Consequently, dinosaur endemism should reflect the past distributions of continents and oceans. One of the conundrums of dinosaur biogeography is the presence of Late Cretaceous dinosaurs in India. According to most plate tectonic models, India was isolated as an island continent for millions of years. Is the concept of India as an isolated island continent compatible with the nature of the fossils found on India ? Current plate tectonic theory postulates that India (Indo-Pakistan subcontinent) together with Madagascar rifted from eastem Africa during Late Jurassic (150 Ma); in the Late Cretaceous it separated from Madagascar and drifted northward 5000 km, finally colliding with Asia to give rise the Himalayas in early Eocene (50 Ma)’. If India were isolated for more than 100 million years, as is usually believed, such an extended period of isolation should have produced a highly endemic Late Cretaceous dinosaur fauna. No such endemism can be seen among Indian dinosaurs. Instead, dominant dinosaurs such as abelisaurs and titanosaurs in India show striking similarities with those of Europe *, Africa’, South America®’, and Madagascar’. If India were separated from adajacent land masses by deep oceanic barriers more than S00 km wide, “Where did Late Cretaceous Indian dinosaurs come from?", and “How did they get there?”. It seems likely that Late Cretaccous India was connected to other lands SANKAR CHATTERIFE AND CHRISTOPHER R SCOTESE during the closing moments of dinosaur evolution, and was not an isolated, island continent. There appears to be a contradiction between dinosaur biogeography and the accepted paleoposition of India at the end of the Cretaceous", To resolve this apparent contradiction, we propose that during the Late Cretaceous a previously unrecognized northem migration corridor opened up between India and Africa. This corridor, through a land arca we term “Greater Somalia” allowed a new influx of dinosaurs from Africa and a homogenization of Gondwana dinosaurs. Fifteen plate tectonic reconstructions are presented illustrating the breakup of Pangea and the cvolution of the Indian platc from its Gondwana origin to its final union with Asia. Using Mesozoic palaeobiogragraphic maps as a framework. the immigration and emigration of dinosaurs and other terrestrial vertebrates in and out India is discussed. Such a synthesis is essential; both geophysical evidence and fossil record must he combined to solve this biogeographic paradox. Paleogeography How Maps Are Made The Mesozoic and Cenozoic plate tectonic reconstructions showing the evolution of the Indian plate and the surrounding oceans since the Late Permian are based on data compiled as part of the PALEOMAP Project of the — International Lithosphere program’. These maps are based on five lines of evidence: paleomagnetism, linear magnetic anomalies, palaenhingengraphy, palaeocli- matology, and geologic history. Although these digital waps are generally based on sea-floor spreading data, the tectonic features of the ocean floor and continents are alsq incorporated using GIS technology. These maps are continuously updated as new evidence becomes available. The Paleo-GIS runs under ArcView 3.0 and produces plate tectonic reconstructions for any period in Earth history fromthe Late Precambrian (750 Ma) to the present-day. ‘Ihese reconstructions have been placed in a reference frame using palaeomagnetic and hotspot data™"”, The 13 time periods covered in this paper showing the palaeocontinental positions of India are: 255 Ma (Longranian. Late Permian), 220 Ma (Norian, Late Triassic), 200 Ma (Sinemurian, Early Jurassic), 180 Ma (Bajocian, Middle Jurassic), 160GONDWANA, EVOLUTION AND BIOGEOGRAPHY OF THE INDIAN PLATE Ma (Oxfordian,. Late Jurassic), 140 Ma (Rerriasian, Farly Cretaceans), 120 Ma (Aptian, Early Cretaceous), 100 Ma (Albian, Early Cretaceous), 90 Ma (Cenomanian, Late Cretaceous), 80 Ma (Campanian, Late Cretaccous, 65 Ma (Cretaceous-Tertiary boundary), 60 Ma (Thanetian, Early Palaeocene), 50 Ma (Ypresian, Eocene), and present-Day. In addition, a biogeographic map showing the possible dispersal routes of Maastrichtian dinosaurs at 66 Ma is also included. With the exception of Fig. 3, these maps show the location of India in orthographic projections. We do not know the original extent of the northern margin of India, because that margin has been extensively deformed by the collision with ‘Asia. One of the important consequences of the Indian plate reconstruction is the recognition that in Gondwana assembly India must have extended farther to the north and east than its present exposure in the southern Himalayas. This enlarged 309 continental fragment, called Greater India’ is necessary to fill the gap between India and the western margin of Australia prior to continental separation at early Cretaceous (130 Ma). Greater India was probably subducted beneath Asia during the initial phases of the collision, and now lies buried in the Tibetan Plateau. The geological time scale discussed in this paper are adopted from Palmer"*. Tectonic Features of the Indian Ocean ‘The Indian Ocean 1s the result of sea-tloor spreading between the African, Antarctic, and Indian plates since the Late Jurassic. Its complex tectonic history has been revealed by geophysical and geological evidence'*"’. The seafloor of the Indian Ocean is dominated morphologically by three active midocean ridge systems that meet at the Rodriguez triple junction (RTJ). They are: the Southwest Indian Ridge (SWIR), the Southeast Indian Ridge (SEIR), and the Central Indian Ridge Fig. 1 Present day plate Map of the Indian Occan illustrating the location of mid-ocean rifts (heavy black line) and submarine plateaus. Abbreviations: BR, Broken Ridge; CH, Chagos; CIR, Central Indian Ridge; CR, Carlsberg Ridge; KE, Kerguelen Plateau; LA, Laccadtve; MA, Madagascar; MAL, Maldive; MTF, Masirah Transform Fault; OER, Ninetycast Ridge; SE, Seychelles.400, (CIR), and its northwestem continuation, the Carlsberg Ridge (CR) (Fig. 1). The SWIR separates Africa and Antarctica and connects with the Mid-Atlantic Ridge. The SEIR separates the Indo-Capricom-Australian plate” from Antarctica and joins with the East Pacific Rise. North of the RTJ, the CIR runs almost due north and turns northward to form the Carlsberg Ridge. One of the striking geological features of the Indian Ocean basin is the large number of long, linear, aseismic ridges and plateaus. These plateaus, as well as several microcontinents, have been proposed as possible dispersal routes for migration of animals to and from India’. The Ninetyeast Ridge (90ER) and the Mascarene Plataeu (MP)- Laccadive (LA)- Maldive (MAL)- Chagos (CH) Island Ridge are two parallel volcanic lineaments that record the northward motion of India over the stationary hotspots at Kerguelen (KE) and Reunion (RE), respectively”. Various continental fragments such as the island of Madagascar, the Seychelles, the Broken Ridge, and the Crozet Plateau became isolated during fragmentation of Gondwana. Gondwana Fit and “Greater Somalia” The fit of the continents that form the core of Gondwana (South America, Africa, Madagascar, India, Australia, and Antarctica) is well known”. Formed during the Pan-African orogeny, approximately 600 Ma, Gondwana remained intact for over 400 My. However, it is interesting to note that in most Gondwana reconstructions there is a gap between northwest India and the southeaster margin of the Indian peninsula. We propose that this gap was fillled by a continental land area which we call Greater Somalia. Though the existence of greater Somalia is required by the geometric fit of the continents, it is not clear what modem land area corresponds to Greater Somalia. One possibility is the Lut block of southeastern Iran (Fig. 1). Traditionally, the Lut block is thought to have arrived at the southern margin of Eurasia during the Late Triassic-Early Jurassic”. However, the Lut block is nearly surrounded by a chaotic assemblage of Late Cretaceous ophiolites (Colored melange), suggesting that it could not have heen incorporated into the southern margin of Eurasia prior to the catly Testiary™. The Eaily Paleozvic SANKAR CHATTERJEE AND CHRISTOPHER R SCOTESE. carbonate sequence of the Lut block are also similar to rocks in Oman and southeastern Arabia’’, and palaeomagnetic data from the Lut block places it in the southern hemisphere, near Arabia during the Devonian’?. Plate tectonic models indicate that India may have collided with Greater Somalia approximately 70-75 Ma (Campanian-Early Maastrichtian). After the collision with India, Greater Somalia slid northward along the transform fault parallel to the coast of southeastern Arabia. Greater Somalia provides a new corridor between India and Africa during Late Cretaceous for migration of dinosaurs and other vertebrates. Whatever its original extent, Greater Somalia was dismembered after the collision with India and may have been partially, or entirely, subducted beneath Eurasia Tectonic Evolution of the Indian Plate The evolution of the Indian plate is the most complex and least understood of all Gondwana landmasses. There are many, reasons for this complexity. One reason is the fact that intraplate deformation has taken place within the Indo- Australian plate. It is now believed that Indo- Australian plate, since its collision with Asia, is made up of three separate plates: India, Australia and Capricorn’. The plate boundaries between them are diffuse and complex extending across a wide zone of deformed lithosphere in the Indian Ocean. A second complexity arises from the fact that a large asteroid impact may have caused the large (2500 km) jump of the Carlsberg Ridge from its location south of the Seychelles Islands, to a new location between India and Seychelles at 65 million ycars ago"*. This episode led to the reorganization of the plate boundaries in the northern Mascarene Basin resulting in the separation of the Seychelles from India. ‘Thirdly, the timing the collision of India with Asia is still debated. Estimates for the age of collision range from 65 Ma to 40 Ma. This collision age is crucial to understand the palaeoposition of India with respect to Asia during the final days of the dinosaurs. India did not rift apart from Pangea in one simple motion but rather its separation from Gondwana ocenrred during several distinct episodes. Ocean magnetic anomaly data indicate eight phases in spreading history of Indian plate. InGONDWANA. EVOLUTION AND BIOGROGR APHY OF THR INDIAN PLATE the next section we describe each of these plate tectonic phases, Chronology of the Evolution of the Indian Plate The dating of the breakup of Pangea and separation of its fragments can be approached several ways. Linear magnetic anomalies and deep sea drilling data provide fairly accurate information about the fragmentation, but there are no anomalies during the long Late Cretaceous, Quiet interval (83-119 Ma). Information from different fields of continental geology and palaeomagnetism also provide additional evidence For example, the eruption of large quantities of ood basalts along the spreading margin may signal the beginning of continental rift. Similarly, marine ansgression associated with cooling and subsidence along the coastal areas may also indicate formation of a new passive margin, There appears to be a long time interval (several tens of millions years) between the tirst indications of continental rift, as shown by volcanism and crustal subsidence, and the subsequent _sea-floor spreading. From a biogeographic point of view, the time of actual continental separation is more g 5 | ve JURASSIC 401 relevant for dispersal and will be used here to discuss the major events of fragmentation of Pangea. A cladistic terminology is proposed here to reflect the evolution of the India plate in a temporal hierarchy as it fragments successively from Pangea, Gondwana, and Indo-Madagascar- Seychelles block, and regroups with Greater Somalia and finally w Asia (Table ). The major nodes of area cladogram depicting the evolution of the Indian plate are shown in Fig. 2. 1 Pangea (255 Ma) During Late Palaeozoic time the Palaeozic continents of Laurentia, Siberia, Baltica and Gondwana moved closer and closer together until by the Early Permian (255 Ma) they were fused together to form the supercontinent Pangea (Fig. 3). Centered on the Equator, Pangea stretched from North Pole to South Polc, and separated the Palaeo-Tethys Ocean to the east, from the Panthalassic Ocean to the west. India was nestled in the Pangea at high southerly latitudes, 2 Gondwana (180 Ma) The Pangea was intact during the Late Triassic Fig. 2 Area cladogram showing the evolution-of the Indian plate from its Pangean origin to the present day.4n2 SANKAR CHATTERJEE AND CHRISTOPHER R SCOTESE. Late Permian 255 Ma Fig. 3 Palucogographic reconstruction of continental positions during the Late Permian (255 ma) showing the supercontinent Pangea surronnded hy the ancestral ocean Panthalassa, Late Triassic’ 220 Ma Fig. 4 Palacogographic reconstruction of Gondwana showing the India’s position during the Late Triassic (220 Ma) period. Note the location of Greater Somalia”, a land afea that was originally north of India. Dinosaurs and other vertebrates were able to migrate freely between all, Gondwana csGONDWANA, EVOLUTION AND BIOGEOGRAPHY OF THE INDIAN PLATE. ysseang aIppryy 24) Bu 1unp uorsoc ‘uortonnisuoza! 31 iydexFoRoseyeg9 “Fy ‘pouad (ep 007)“ S.eIpUL a4 Buywoys eseMpUoN Jo —<—o issvang Kye ay uuunp uantsod wonoansuonar onpleiBoFozetd ¢ “RyTable T Evolution ofthe Indian plate during the Permian Mesozoic ‘and Cenezoic Perieds associated with plate tectanic events Continental Lan ents masses Containing India 1 Pangea Formation of the supercontinent Pangea with Paleo-Tethys (255 Ma) 2 Gondwana Rift in Pangea to form Condwana and Laurasia (180 Ma) 3 East Gondwana Rift Between West and East Gondwana (160 Ma) 4 Indo-Madagascar-Indo-Madagascar-Seychelles separates Seychelles with Antarctico-Australia (12 Ma) 5 Indo-Madagascar- Brief isolation of Indo-Madagascar- Seychelles, island Seychelles (100 Ma) continent © Indo-Sechelles, island continent 7 Indo-Seyehelles- Greater Somalia Indo-Seychelles separates from Maula gascar (RR Ma), brief isolation Collision of lndu-Seyelelles with Greater Somalia (70 Ma) contacts with Africa 8 Indo-Greater Indo-Greater Somalaria separates from Somalia Seychelles (65 Ma); Shiva impact Y India India separates from Greater Somalta (60 Ma) Collision of hadi wide Asia, disap= pearance of Tethys (51 Ma) India sutured to Asia; subduction of Greater India; rise of the Himalayas (20 Ma) 10 India contacts Asia 11 Ludu-Asia and Barly Jurassic periods (Figs 4 and 5). The first stage of rifting of Pangea took place in Middle Jurassic period (180 Ma) between North America and Africa after an episode of volcanic activity, leading to the separation of Gondwana and Laurasia (Fig. 6). At that time India was still part of Gondwana. 3 East Gondwana (160 Ma) During the Late Jurassic, Gondwana began to rift apart. The initial phase of rifting, was signalled by the eruption of widespread flood basalts, such as Karoo Group in South Africa and Ferrar Group in East Antarctica. During the first phase of continental rifting Gondwana broke apart into two roughly equal parts: western Gondwana composed of Africa and South America, and easter Gondwana composed of Madagascar, India, Australia and Antarctica, The oldest ocean floor separating the western Gondwana from eastern Gondwana is Late Jurassic, approximately 160 My old (Fig. 7). This phase in the breakup of Gondwana extended thuugh Jurassic intu he SANKAR CHATTERJEE AND CHRISTOPHER R SCOTESE earliest Cretaccous (Fig. 8, 140 Ma) resulted in the formation of Somali Basin (SB) and Mozambique Basins (MB) (Figs. 7 and 8). 4 Indo-Madagascar-Seychelles (120 Ma) During the Early Cretaccous (120 Ma), both the western and eastern halves of Gondwana began to split in two (Fig. 9). South America hegan to separate from Africa forming the South Atlantic, and Indo-Seychelles-Madagasca_bluck separated from Antarctica and Australia opening the Central Indian Ocean. Both of these oceans opened progressively. The South Atlantic opened from south to north, and the central Indian Ocean opened from northeast southwest (Fig. 9). By the mid-Cretaceous, sea floor spreading in the Somali basin had stopped. Indo-Madagascar- Seychelles block hecame fixed to African plate, 5 Indo -Madagascar-Seychelles Island Continent (100 Ma) Sea-floor spreading between Indo-Madagascar- Seychelles block and Antarcto-Australia continued widening the Central Indian Ocean so that the former landmass became completely isolated during the later phase of the Early Cretaceous (100 Ma) and completely surrounded by the ocean (Fig. 10), The South Atlantic also continued to widen as Africa and South America became completely separate continents. In the south central Indian Ocean, an extensive volcanic plateau, Kerguelen was generated at the continuation of the Rajmahal hotspot of India that would create the 90 East Ridge”. During this time, Kerguelen plateau was subaerial and may have provided an interminent dispersal route between India and Antarctica’. 6 Indo-Seychelles Island Continent (90 Ma) During the Late Cretaceous (YU Ma), Indo- Seychelles rifted away from Madagascar to open the Mascarene Basin (Fig. 11). Indo-Seychelles began to move rapidly northward (17cmvyr), and by 80 Ma (Fig. 12), it was completely separated from the other Gondwana continents. 7 Indo-Seychelles-Greater Somalia (70 Ma) Sometimes during the latest Cretaceous (70 Ma), India collided with Greater Somalia to form a biogeographic corridor with Africa. This collision ended India's brief period of isolation (Fig. 13). AtLate Jurassic 160 Ma ge GONDWANA, FVOLLTION AND BIOGEOGRAPHY OF THE INDIAN PLATE 405SANKAR CHATTERJEE AND CHRISTOPHER R SCOTESE JOpUsca WAKIM AH, Hopitios woy;nog =g s1epis409 BW OZ} Snosceelg AYESGONDWANA, EVOLUTION AND BIOGEOGRAPHY OF THE INDIAN PLATE “suounuoo euempuog 210 -opyy ‘poucd (eN Og} smi BURAPIOD J suoasod ay ButaSANKAR CHATTERJEE AND CHRISTOPHER R SCOTESE, ey $9 Arepunog iseye-snoeoeies)GONDWANA. EVOLUTION AND RIOGEOGRAPHY OF THE INDIAN PLATE, about the same time, dispersal routes between Europe and Africa, between Europe and North America, and between North and South Americas were reestablished. These biotic links would eventually connect India with other Gondwana continents. In the Tate Cretaceans, continned spreading along the South East Indian Ridge (SEIR) widened the Indian Ocean between India and Antarctica making Ninety East Ridge ineffective for biotic disperal between these two landmasses. 8 Indo-Greater Somalia (65 Ma) A new rift between Indo-Somalia and Seychelles was formed near the KT boundary (65 Ma), coinciding with the dinosaur extinction (Fig. 14). At this time, the Central Indian Ridge (CIR) jumped 500 km northward from its location in the Madagascar Basin to a new location between Seychelles and India (the Carlsberg Ridge, CR). This event resulted in sudden separation of Seychelles from -Indo-Greater Somalia (Fig. 13) and the transfer of the Seychelles and Mascarene bank to the African plate. Seychelles may have behaved as an independent microplate during this rifting event. This ridge jump (> 500 km) may be linked to the Shiva impact on the trailing edge of the Indo-Seychelles block”, This impact may have triggered readjustments along the Indian- African and Antarctic-African plate boundaries by causing lithospheric weakness that would have helped to create the Owen Fracture Zone (OFZ) and Carlsberg Ridge”. 9 India (60 Ma) India continued its rapid approach towards Asia. During the Paleocene (60 Ma), Greater Somalia was carried northwards with India, parallel to the coastline of Arabia, and moved towards Iran- Afganistan (Fig. 15). Much of Greater Somalia may have been subducted during Miocene collision beneath Asia that formed the Makran ranges. 10 India-Asia initial contact (50 Ma) The timing of collision between India and Asia has been subject of much debate. Estimates range from the Late Cretaceous (>65 Ma)” to latest Eocene (<40 Ma)’. The age of the collision is based primarily on a change in the speed of the Indian plate. In the Palacocene, India slowed its 409 northward motion from about 20 cmv/year to 4.5 em/year, indicating its suturing with previously accreted terranes; direct subaerial connection with Asia probably occurred later. Recent compilation of data from linear magnetic anomalies in the Indian Ocean as well as palacomagnetic, voleanic eruption, and stratigraphic evidence from the Indian subcontinent indicate that the initial collision took place between 55 and 50 Ma in Early Eocenc™ along the NE corner of Greater India. The initial collision marks the cessation of marine sedimentation and the commencement of terrestrial deposition in the suture zone, allowing a subaerial link between India and Asia (Fig. 16). 1 Indo-Asia (20 Ma) Since initial collision: with Asia, India has moved northwards an additional 2000 km, resulting in spectacular rise of the Himalayas. Further convergence has led to gigantic east-west ending strike-slip faults that are forcing eastern Asia to slide to the east, out of India's path’. The rise of the Himalayas took place much after the time. of intitial collision of India with Asia by the horizontal compression and subduction of the continental lithosphere (Fig. 1). One effect of the Himalayan orogeny is construction of the Bengal Fan, the world’s largest submarine fan, with sediments derived from the uplift and erosion of the High Himalayan crystallines. Sedimentation in the Bengal Fan delta provide additional evidence for the timing of the Himalayan uplift. The oldest turbidites deposited offshore from the Indus and _Ganga rivers date ack to the Lower Miocene (17 Ma), which pushes back the main phase of the Himalayan uplift around 20 Ma™. Indeed much of the Himalayan chain has been uplifted during the last 15 My. Although India is now geographically an integral part of Asia, Himalayas form both orographic and climatic barriers for dispersal of terrestrial vertebrates between these two areas. Biogeography of Dinosaurs Dinosaurs are the most spectacular and captivating animals known to human, representing the glory of evolution and mystery of extinction. They appeared in the Late Triassic (227Ma), evolved into some of the largest and most successful land animals of all time during Jurassic and Cretaceous periods, and then suddenly disappeared at the endSANKAR CHATTERJEE AND CHRISTOPHER K SCOTESE the Paleocene (60 Ma) period. India pr corndor through Greater Somalia. AtbreGONDWANA, EVOLUTION AND RIOGFOGRAPHY OF THE INDIAN PLATE of the Cretaceous (65 Ma). For 162 million years, they ruled the Earth and flourished on every continent including Antarctica. They adapted to nearly every evironments--deserts, hot lakeshores, volcanic provinces, sandy beaches, swamps, tropical and temperate jungles, and even polar forests. When considering dinosaur biogeography, several factors should be taken into account. First, the fossil record of dinosaurs is poorly represented, and many taxa are known from incomplete material. There are currently 325 valid genera of dinosaurs, which may represent about 8%” to 30%" of the total number of genera that may have existed. Second, there are sampling biases. Many continents are not thoroughly explored for dinosaur bones. In fact, because of resurgence in dinosaur research, 50% of new dinosaur genera have been discovered since 1970, many of them from exotic lands such as China, Mongolia, Argentina and Antarctica, and many more areas still need to be explored. Third, their temporal distribution is uneven. For example, 30% of dinosaur genera are known from the Late Cretaceous” alone, whereas Triassic and Early Jurassic records arc relatively sparse. In spite of these deficiencies, an interesting pattem of dinosaur distributions 1s slowly emerging in the context of plate tectonics. Biogeographic Principles Biogeography attempts to explain the distributions of organisms in terms of climatic and geographic barriers to migration. Two principal factors control the distribution of land animals: climate and sea barriers. Climate may not be a limiting factor for the distributions of dinosaurs, because their fossils have been found near the poles, in the temperate regions, in the tropics, and in deserts. They did not habitually Swim and never acquired aquatic adaptations. For dinosaurs, the yea way a fouinidable barrier whether it be shallow or deep. As a result, they are likely to provide a better guide to the chronology of separation and fusion of continental fragments during the Mesozoic than plants and __ invertebrates. Occasionally dinosaur bones have been found in near-shore marine environments, The taphonomy of these occurrences suggest that floating carcasses aul or bones were transported into the marine burial site hy rivers In order to understand the global distribution of dinosaurs during the Mesozoic, two common principles of biogeography, namely vicariance and dispersal should be discussed. The respective role of vicariance and dispersal in controlling the distribution of organisms has been debated vigorously. Vicariance biogeography” considers the history of distribution of organisms in relation to the history of the plate movements. It requires the formation of barriers after the organisms have established their early biogeographic range. Hallam™ discussed two models of biogeography of land vertebrates within the vicariance backdrop. When a continent splits, the cosmopolitan populations are isolated from each other leading to genetic isolation and morphologic divergence. Conversely, when two continental fragements collide, there is cross migration leading to more uniform and homogeneous tauna. Both phenomena may be true for India during the Cretaceous period. Dispersal biogeography, on the other hand, was developed during the time of Darwin on the premise of fixed continents with numerus hypothetical landbridges. In this model. the main mechanism of, distribution of organisms across continents and scas is their own dispersal, whether by walking, flying, swimming, or even by rafting. Simpson” recognized three types of dispersal routes for land animals: corridors, involving land connections large cnough to allow free movements of animals in both directions; filter bridges, which involve land connection with other factors such as climate or mountain that function as barriers to some elements of a biota but not to others; and sweepstake routes, along which dispersal involves a large element of chance like rafting across a sea and colonizing in islands. In dispersal view, the center of origin of a clade is an important issue. ‘The difference between the two schools of biogeography, as Molnar” succintly pointed out is that vicatiance bivgeugraphy postulates that continents move, but not organisms, whereas dispersal biogeography assumes that organisms move, but not continents. In reality, hoth vicariance and dispersal are important Zomponents in biogeography, especially for the distribution of dinosaurs. Dinosaurs lived on all seven continents. Because412 of shifting continental positions and associated rifting and suturing events, vicariance, biogeography must have played an important role in the distribution of dinosaurs in time and space. However, the role of dispersal cannot be ignored in, dinosaur biogeography. All large dinosaurs, herbivores and predators alike, would have to make long migrations in order to gather enough food to keep themselves going. Dinosaurs were thorough-going terrestrial anaimals. Because of their erect posture and refined locomotion, they were “happy wanderers” and many large dinosaurs were capable of long-range seasonal migrations similar to modem African animals®®. The covered by dinosaurs would be close to about 30° latitudes (~3,000 km) in each way during food gathering season, indicating that they were widely dispersed within a continent. Indian Dinosaurs The first dinosaur remains to be described from Asia were found in India. Richard Lydekker” trom the Geological Survey of India (GSD) described a new genus and species of a sauropod dinosaur, Titanosaurus indicus from the Late Cretaceous Lameta formation of Central India. Since then, however, Indian dinosaurs. which are often represented by incomplete specimens, have been somewhat overshadowed by spectacular finds of Tal Terrestrial tetrapod-bea SANKAR CHATTERJEE AND CHRISTOPHER R SCOTESE complete skeletons and diverse fauna from China and Mongolia Dinnsanr hones have heen collected sporadically by the GSI from the Cretaceous beds since Lydekker’s report, and the whole collection was described in a monograph including fragmentary remains of sauropods, theropods and an ankylosaur™. Systematic collections of dinosaur bones from different formations of Gondwana sediments began in 1960s by the Indian Statistical Institute (IST), followed by the Geological Survey of India and the Panjab University (PU). Dinosaur-Bearing Gondwana Formations ‘Assessment of the palaeopositioin of India during the Mesozoic depends heavily upon the distribution of terrestrial tetrapods. This study focuses on dinosaurs in general, as well as contemporary terrestrial tetrapods, excluding the volant forms such as pterosaurs and birds. For flying animals, the sea may not be a barrier for dispersal. In view of the limited dinosaurian record, we have also included other terrestrial tetrapods for faunal correlation between India and other continents. India has one of the finest uninterrupted Gondwana sequences spanning from Early Permian to Late Cretaceous, and should preserve a continuous history of the rise and fall of dinosaurs. In reality, India is scarce in dinosaur data. There are several factors for this limited ble It iring Gondwana formations (indicated by an asterisk) MA PERIODS FORMATION IMPORTANT TETRAPOD FAMILIES 65. 8 Titanosauridae, Abelisauridac, Nodosauridae, Bh LAMeTA 7 on BP LAMETAM*ARIYALUR/ petomedusidae, Pisenglossidne, Palaedryctidae 100- Q _PINTERTRAPPEANS 120- 3 IMAR 140 9 |cuIKIALAGANGAPUR 160- BAGRAIPATCHAM Diplodocidac, Stegosauridae Q | 180- @ — (RAJMAHAL 200- 5 bKora Vulcanodontidae, Teleosauridae, Kuehneotheridae 220 | DHARMARAM Plateosauridae, Thecodontosauridae, Saltoposuchidae, Q Herrerasauridae, Phytosauridae, Rauisuchidae, Pro- % lacertidae, Rhynchosauridae, Traversodontidae, Me~ 240- SP MALERI toposauridae, ChigutisauridaeGONDWANA, EVOLUTION AND BIOGEOGRAPHY OF THE INDIAN PLATE knowledge of Indian dinosaurs. First, many areas and several potential horizons have never been throroughly prospected. Second, the land is densely populated and intensely cultivated, thus destroying many paleontological sites. Third, there is a direct correlation between the habitat area and the diversity of terrestrial species: the larger the area, the higher the diversity of species. India, being one of the smallest landmass among Gondwana continents, would support few species of larger dinosaurs after fragmentation. In spite of these deficiencies, dinosaur research is gaining currency in India in recent times. The finest collections of Indian dinosaurs and their contemporaries are housed at the Geology Museum of the Indian Statistical Institute, Calcutta. Other collections can be seen at the Geological Survey of India at Calcutta, Gandhi Nagar, and Jaipur. Panjab University at Chandigarh possesses a large and important collection microverbrates associated with dinosaurs. Dinosaur-bearing horizons in Indian Gondwana formations are shown in Table IL 413 As an index of faunal similarity between two regions, Simpson” suggested the simple formula of 1OOC/N 1, where C is the number of faunal units of given taxonomic rank, common to two areas, and N1 is the total number of such units in the smaller of the two faunas, In our discussion, NI represents the total number of tetrapod taxa present in India. ‘The advantage of this formula is to minimize the effect of unequal size of the faunas being compared. The faunal comparisons are made at the family level. The classification of dinosaurs, and stratigraphical distributions of families are taken from Weishampel et al.”’. J. Late Triassic Vertebrates (Carnian, 227 Ma) The Late Triassic Period was an extraordinary time in vertebrate evolution when several major groups such as dinosaurs, birds, crocodilians, squamates, lissamphibians and mammals first appeared in the fossil record. This faunal diversification heralded the beginnings of many major groups of tetrapods that still persist today. Since the Pangea was symmetrically disposed ‘Metoposauras (right) and Compsocerops (left). In the foreground left is the rauisuchid Tikisuchus, center herrerasaurid Alwalkeria, and right is the thynchosaur Paradapedon.414 above and below the equator, the topography and climate became more uniform. The paleoclimate was dominated by monsoonal circulation. Dense forest and swamps covered the land from the equator to high latitudes. In this Pangean world, terrestrial vertebrates were able to migrate easily back and forth across the land surface of the Earth Fig. 4). Dinosaurs—One of the earliest dinosaurs, Alwalkeria is known from the Maleri Formation of Godavari Valley. It was a small theropod, about a meter long. Current study suggests that it 1s very similar to Herrerasaurus of Argentina and Chindisaurus of Arizona (Fig. 17). Other Tetrapods—Although Alwalkeria is the sole dinosaur so far known from the early Late Triassic (Carnian), several tetrapods trom the Maleri Formation suggest that India was the crossroad of faunal migration between Laurasia and Gondwana. For example, travesodont Exaeretodon® rauisuchid —Tikisuchus and thynchosant Paradapedon™ are very similar to those of Europe, North America, Africa and South America. Ou the vthe: hand, phytusaurs and ‘tmetoposaurs from the Maleri Formation are comparable to those of North America Europe, Morocco and Madagascar, but are cospicuously absent in South America“*. Both were semi-aquatic animals like modem crocodilians and were probably controlled by latitudinal variation. Their absence in South America, Australia, and Antarctica may be associated with relative paleolatitudinal position in Southern Hemisphere below 30°S (Fig. 4). The prolacerid Malerisaurus has also been identified in North America®, SANKAR CHATTERJEE AND CHRISTOPHER R SCOTESE Sengupta’’ described two genera of chigutisaurid amphibians, Compsacerops and Kuttycephalus from the Maleri Formation. The chigutisaurids are known from the coeval beds of South America and recently from North America. It appears from Table I that the Maleri fauna is overwhelmingly ‘northern’ and show the strong correlation with Africa and North America. Possibly. the route of faunal migration between India and North America was via Morocco, as similar taxa have been found there. On the other hand, Indian tetrapods are not represented in Australia and Antarctica possibly because of lack of coeval beds. 2, Late Triassic Vertebrates (Upper Norian, 215 Ma) Vertebrates from this age are known from the Dharamaram Formation of Godavari valley. The presence of two prosauropod dinosaurs, an omithischian, and a sphcnosuchian crocodylo- morph from this horizon has been briefly reported three decades ago“. The material is beautifully preserved, but lack of description and illustration makes the comparison difficult. Dinosaurs—The prosauropod fauna includes a large plateosaurid and a small thecodontosaurid (Fig. 18). The plateosaurid is represented by an associated skeleton of a fairly complete postcranial skeleton as well as few cranial fragments. In size and proportion, it is very similar to Plateosaurus of Germany. The thecondontosaurid contains a partial vertebral column and limb elements. It compares well with Thecodontosaurus of Great Britain. The omithischian is represented by pelvic and sacral clements, but its affinity remains unknown. Table 111 Indices of Faunal similarities of Indian Tetrapod families during the late Triassic (Carnian) period Indian Genera Andtamilies ‘Europe —N.Amenca Asia Atica_—‘S. America Australia Antarctica ‘Abwalkeria, — Heerrerasauridae = + = + 7 = a Parasuchas: Phytosauridac + + . + 5 6 - Tikisuchus: — Ravisuchidae - - - + + Malerisaurus: Prolacertidse e + - Paradapedon: Rhynchosauridae + + 5 + + - 2 Exaeretodon: Traversodontidac + + - + + 5 S Metoposaurus: Metoposauridae + + 5 + e e . ‘Compsocerops, Chigutisauridae - + - 5 + 5 - Kuttycephatus Index 30 88 0 5 50 0 0GONDWANA, EVOLUTION AND BIOGEOGRAPHY OF THE INDIAN PLATE 45 Fig. 18 Upper Triassic (Norian) Dharamaram vertebrates. In the foreground is the larger prosauropod, plateosaur. In the background is the smaller prosauropod thecodontosaur. Between the hindlimbs of the plateosaur, a sphenosuchid creodylomorph can be seen, Table1V Indices of Faunal similarities of Indian Teirapod families during the late Triassic (Norian) period Indian Genera Andfamilies Europe N. America Asia_—_—Affica.—S. America Australia Antarctica Unnamed Plateosauridac + = 5 = + 5 = Unnamed ——Thecodontosauridae + - - . = = - Unnamed __ Saltoposuchidae + 2 = = = S| = Tex 100 0 0 3 Other Tetrapods—The sphenosuchian material shows articulated shoulder girdle, forelimb and anterior dorsal vertebrae and is reminiscent of Saltoposuchus of Germany and Wales. Although the Dharmaram fauna is incompletely known at present, it shows the highest correlation with European fauna (Table IV). The lack of correlation with other continents (other than Europe and South America), may be an artifact; Upper Norian continental sediments containing vertebrates are rare. 3. Early Jurassic Vertebrates (Toarcian, 200 Ma) The end-Triassic extinction led to! the disappearance of several well-known groups of nondinosaurian tetrapods including metoposaurs, phytosaurs, stagonolepidids and procolophonids. Dinosaurs survived this catastrophe and rebounded with new vigor. Pangea was still intact during the Early Jurassic period allowing dispersal of dinosaurs unhindered across the globe (Fig. 5). Dinosaurs—Iin India, the Kota Formation of Godavari Valley has yielded spectacular skeletons of early sauropods such as Barapasaurus”. The skeleton is gigantic in the context of its early geologic occurrence. It is about 16 m long and 4m high, and has developed all the sauropod attributes. Barapasaurus is comparable with Vulcanodon of416 Zimbabwe, Ohmdenosaurus of Germany and Zichonghosaurus of China (Fig. 19). An ilium at the GSI collection from the Kota Formation indicates the presence of an omithischian dinosaur’, Other Vertebrates—A teleosaurid crocodilian has been beautifully prepared at ISI and awaits description. Similar crocodilian has been reported from Europe, North America, Africa, and South America. The early Jurassic mammals of Kota Formation include several genera of kuehneotheriids including Kotatherium®' and Trishulotherium™. This group of early mammal is so far restricted to the coeval beds of Europe. 2 dt GE SANKAR CHATTERJEE AND CHRISTOPHER R SCOTESE ‘The faunal correlation between India and Europe is highest during early Jurassic period (Table V). 4, Middle Jurassic Vertebrates (Dogger, 178-157 Ma) The fossil record of Middle Jurassic dinosaurs are rare, Scattered localities are known from Europe, Asia, South America, Africa and Australia. Russell suggested that China became isolated from the rest of the Pangea from Middle Jurassic into Early Cretaceous periods and evolved many endemic forms. During the Middle Jurassic, Gondwana began to split into western and eastern components, but India remained in substantial Fig. 19 Lower Jurassic Kota Vertebrates. In the foreground is the teleosaurid crocodile. In the background, one of the earliest sauropod dinosaurs Barapasaurus. Table V Indices of Faunal similarities of Indian Tetrapod families during the early Triassic (Norian) period Indian Genera And families Barapasaurus. Vuleanodontidae Burope _N. America + 7 Asia Africa 8. America Australia Antarctica = + = = = Unnamed ——Teleosauridae + + = + S S 7 Kotatherium: Kuehneotheriidae + = = = = - Trishulotherium Tndex 100 3 0 66 33 0 aGONDWANA, EVOLUTION AND BIOGEOGRAPHY OF THE INDIAN PI.ATF. contacts with both components allowing faunal interchange (Fig. 6). Predicting the ‘Undiscovered _Indian Dinosaurs—Middle Jurassic dinosaurs are unknown from India, but we can make predictions about the expected dinosaur faunal composition from the plate tectonic reconstruction (Fig. 6) and the distribution of fauna in other parts of the world adjacent to India (Table V1). During this time, 7 familics of dinosaurs are known from Europc, 4 from Australia, 3 from South America, 4 from Africa, and none from Antarctica. It appears from the Table VI that cetiosaurids, brachiosaurids and allosaurids had a wide range of distribution and may be expected in India during this time if potential beds are found. 5. Late Jurassic Vertebrates (Malm. 157-145 Ma) During the Late Jurassic, Laurasia began to separate from Gondwana, creating an cquatorial seaway between them, leaving great deposits of evaporites along the Gulf of Mexica, and forming, much of the coastline bordering the Tethys sea. Global temperatures were generally warmer Uran today, allowing dinosaurs and other vertebrates to live at higher latitudes. Laurasia was fragmented and Europe became an archipelago. The similarity of dinosaur fauna between Morrison Formation of North America and Tendaguru Formation of eastern Africa indicates that European archipelago 417 allowed limited faunal interchange between Africa and North America until the Late Jurassic. The dinosaur is dominated by a cosmopolitan sauropod-stegosaur-iguanodontian fauna. During this period, India maintained connections with both West and East Gondwana allowing faunal interchange (Fig. 7). Dinosaurs—The Late Jurassic dinosaurs have not been reported from India, but recent tantalizing finds suggest that the continental beds of this age should be thoroughly explored. A small diplodocid sauropod in the collection ot ISI was recovered from the Bagra Formation of Satpura Valley. Similarly, few tail spikes of a stegosaurid omithischain at the collection of GSI were found in the shallow marine Patcham Beds (Callovian) of Kutch (Fig. 20). Diplodocids are known trom Africa, North America, Europe; whereas as stegosaurids have been reported from Africa, North America, Europe, and China (Table VI). 6. Early Cretaceous (146-97 Ma) During the Early Cretaceous, Laurasia and Gondwana fragmented; shallow scas invaded North America and Europe. However, North ‘American and European dinosaurs were very similar during this period, dominated by brachiosaurid-iguanodontid-nodosaur fauna; the similarity indicates land connections. Similar biotic links are known between Africa, Europe and Table VI Tetrapod families during the Middte jurassic pertod tn continents around india Dinosaur families Atica S. America Australia Europe Cefiosauridae + + = Fe Diplodocidae - - - + Brachiosauridae + ' S ' Iguanodontidae + + Stegosauridae + = = + Hypsilophodontidae - - + + Allosauridae + + + + Table VIE Indices of Faunal similarities of indian Tetrapod families during the late Jurassic (Kimmeridgian) period Indian Genera Andfamilics Europe N.America Asia _—Africa_—S, America Australia Antarctica Unnamed Diplodocidae + + = + = 7 Unnamed —_Stegosauridae + + + + i - - Tndex 100 100 30 100 0 0 0418 SANKAR CHATTERJEE AND CHRISTOPHER R SCOTESE Fig. 20 Upper Jurassic Dinosaurs such as stegosaurs (foreground) and diplodocid sauropod (background) lived in India, South America as indicated by the presence of Gondwana elements such as _abelisaurids, titanosaurids and spinosaurids. The occurrence of Gondwanan elements in Europe throughout the Cretaceous is interesting, The sauropods remained a major faunal component in both Laurasia and Gondwana. India became separated from Greater Somalia and Africa, thus closing the northern corridor. However, three other biogeographic corridors, namely central (via Madagascar), Southern (via South Africa/Falklands) and western (via Antarctica) remained open which would allow immigration and emigration of dinosaurs and other vertebrates (Fig. 8, 140 Ma). In India, fragmentary dinosaur bones have been encountered in the Early Cretaceous Chikiala and Gangapur Formations of the Godavari Valley, but their affinity is unknown. Future explorations from these horizons may fill important gaps in the temporal distribution of Indian dinosaurs. Around 120 Ma (Aptian), further fragmentation of Gondwana took place when South America separated from Africa, and India from Antarctica- Australia. However India retained three biogeographic corridors, namely western, central and southem through intermittent connections (Fig. 9). Around 100 Ma (Albian), India together with Madagascar shows complete isolation, but central and western corridors may allow limited faunal interchange by sweepstake routes (Fig. 10). The lack of fossils during the long span of Early Cretaceous makes it difficult to determine the degree of endemism among Indian vertebrates during this isolation period. Predicting the ‘Undiscovered _Indian Dinosaurs—Although the fossil record of Early Cretaceous dinosaurs is unknown from India, we can make predictions what kinds of animals lived there from the continental positions of India in relation to other Gondwana continents. During the early phase of Barly Cretaceous, India had three open corridors (Fig. 9) for faunal interchange. During this time, 10 families of dinosaurs are known from Africa, 4 from South America, 5 from ‘Australia, 10 from Europe, and none from Antarctica (Table VII). The occurrence of wideGONDWANA, EVOLUTION AND BIOGEOGRAPHY OF THE INDIAN PLATE 419 Table VIII Tetrapod families during the Middle jurassic period in continents around India Dinosaur families Attica ‘S. America Australia Europe Brachiosauridae Comarasauridae Dicracosauridae Titanosauridae Carcharodontosauri- dae Elaphrosauridae Spinosauridae Dryosauridae Iguanodontidae ‘Stegosauridae Hypsilophodontidae - Nodosauridae . Allosauridae + - + + + I es + tee ee 1 ' betas mre 1 tee teen Paar right abelisaurid theropod Indosuchus. In the background two titanosaurids, the medium-sized, armored Saltasaurus, and the large Titanosaurus. In the foreground water, pelobatid frog and discoglossid lizard can be seen.420 range of dinosaur families in Europe, Australia, and Africa indicate that some of the common faunal elements such as_brachiosaurids, titanosaurids, iguanodontids, nodosaurs, and allosaurids may have been present in India during the Early Cretaceous period. 7. Late Cretaceous (Maastrichtian 66 Ma) During the Late Cretaceous, the continents continued to separate and began to assume their madern positions. Great warm seas covered much of the Earth during this period, a time ot mild climate and ice-free polar regions. The dispersal of continents separated dinosaur communities into distinct Asiamerican (hadrosaurs, ceratopsids, and tyrannosaurids) and Euro-Gondwana (titanosaurids and abelisaurs) groups“, but intermittent connections allowed sporadic migrations. During the very Late Cretaceous, South America was connected to North America across the Caribbean volcanic chain allowing migration of Gondwanan titanosaurs to North America. India-Madagascar separated from Africa during the Tithonian (Fig. 8, 140 Ma), aud Madagascar separated from India during Campanian (Fig. 11, 90 Ma). India became briefly isolated during this period retaining only the central biogeographic corridor (Figs. 10-12). When India collided with Greater Somalia around 70 Ma, a northern migration corridor with Africa was reestablished (Fig. 13). It is likely that Maastrichtian vertebrates of India reflect a faunal immigration event after the collision of India with Greater Somalia. The collision allowed African and European vertebrates to sweep into India. Dinosawrs—In India, the Laweta Formation, sandwiched between the flows of the Deccan Traps, has yielded one of the finest dinosaur localities. The older collection, described by Huene and Matley™, is housed at GSI, Calcutta. The fauna contains several titanosaurid genera such as Titanosaurus and Antarctosaurus, large theropods such as Indosuchus and Indosaurus, several small and fragmentary remains of theropods of uncertain affinities including Jubbulpuria, Composuchus and Ornithomimoides, and probably an ankylosaur. Chatterjec™ described a partial skull of Indosuchus at the collection of the American Museum of Natural History. An extensive collection of Maastrichtian dinosaurs has been made recently by Chatterjee & Rudra® and is SANKAR CHATTERIFF AND CHRISTOPHER R SCOTESE now housed at ISI. These authors provided the preliminary assessment of the systematic positions of Indian Maastrichtian dinosaurs. They placed Indosuchus and Indosaurus within Abelisauridae which are well known from Argentina, Madagascar, Europe and Africa. Among titano- saurids, both Titanosaurus and Antarctosaurus are represented by new material of braincase and postcranial clements. In addition, a medium-sized armored titanosaur, very similar to Saltasaurus of Argentina is also present in the collection. Titanosaurids had a wide distribution during the Maastrichtian period, known from all continents except Australia and Antarctica, Chatterjee and Rudra collected postcranial elements of a nodosaurid ankylosaur. Nodosaurids were widely distributed during this time including Antarctica (Fig, 21). Other Vertebrates—The palacoryctid mammal Deccanolestes has been reported from the intertrappean beds of Naskal of Godavari Valley”. The genus shows morphological similarity to the North American, European, Mongolian and possibly South American forms. The palacoryetid mammals are also known from the Palaeocene of Morocco, possibly a late arrival from India. Other vertebrates from Maastrichtian beds include a pelumodusid title Shwebverys””, a pelobatid frog, and a discoglossid frog™. None of the vertebrates show any evidence of endemism. The affinity of Indian Maastrichtian vertebrates is shown in Table IX. Again, the index of faunal similarity is the highest between India and Europe. Rouieg of Dispersal of Late Cretaceous Indian Dinosaurs Hallam” cited the distribution of Late Cretaceous dinosaurs of India (such as abelisaurs and titanosaurs) as a prime example of disjunct endemism. He argued that similar dinosaurs are known from South America, Europe, Africa, and Madagascar which were separated from India during the Late Cretaceous time by deep ocean. Since India was isolated for 100 million years. the similariry of dinosaurs in all these areas cannot be explained unlcss India maintained some land connections with other areas in the Late Cretaceons time. Sampson et ai.” addressed this unusual disteittution of abcfisaurs in disjunct JandmassesGONDWANA, EVOLUTION AND BIOGEOGRAPHY OF THE INDIAN PLATE such as India, Madagascar and South America during the Late Cretaceous time. They explored two possible routes of dispersal: one is through northern corridor where abelisaurs migrated to India and Madagascar from Africa sometimes in Early Cretaceous before the major continental fragmentation. The alternate hypothesis suggests that abelisaurs migrated to India and Madagascar 421 during the Maastrichtian period from South America through westem corridor via Antastetica across the Kerguelen Plateau (Fig. 22). We prefer the first hypothesis because of two reasons: (1) Throughout the Mesozoic, India shows the least biotic correlation with Australia and Antarctica (see Tables II-LX). This may be attributable to poor sampling. However, there may be some kind Table IX Indices of Faunal similarities of Indian Tetrapod families during the Late Cretaceous (masstrichtian) period Indian Genera And families Furope N.America Asia _—-Africa_—«S. America Australia Antarctica (Madagascar) Titanosaurus: ‘Vitanosauridae = = 5 3) + 5 = Antarctosau- rusSaltasaurus Indosuchus; — Abelisauridae + . 5 ++) + - - Indosaurus Unnamed: Nodosauridae + + . +) . . + Shweboemys — Pelomedusidae + + + +) a = - Unnamed _—Pelobatidae + + a 4+) S = = Unnamed: Discoglossidae + + 5 + - e = Deccanolestes: Palacoryetidac + + + +2) + e _ index 100 % 3B 3787) a 0 a BIOGEOGRAPHIC CONNECTIONS TO INDIA Ss c N NE Fig. 22. Biogeographic connections to India during the last 200 million years showing the fate of various corridors. Abbreviations: C= Central corridor; N= Northen corridor; NE= Northeastem corridor; S= Southern corridor; W= Western corridor.422 of ecological barrier between India and Australia- Antarctica. AS a result, the migration from Amlarctica to India is unlikely. (2) By Maasatrichtian time, the Kerguelen Plateau appears to be separated from India by a vast stretches of ocean so its role as a possible land bridge between India and Antarctica is doubtful. The migration of dinosaurs and other vertebrates from Africa to India through the Greater Somalia appears to be more parsimonious (Fig.13). We get a better picture of biotic dispersal route when we consider the affinity of faunal assemblages instead of a single clade of dinosaurs (Table IX). We see the highest faunal correlation between India and Europe. The route of faunal dispersal must be through Greater Somalia to India along northern corridor. The moderate correlation between India and Africa may reflect poor sampling. On the other hand, the earliest representatives of titanosaurs and abelisaurids are known from Africa. Janenschia is the oldest titanosaurs known from the Upper Jurassic sediments of Africa, succeeded by the Middle Malawisaurus”', Similarly, the earliest abelisaurid Majungatholus is known from the Middle Cretaceans of Moraceot speculate that ancestors of both titanosaurids and abelisaurids lived in Africa before the major breakup of Gondwana and spread later to South ‘America, Indo-Madagascar and Europe. This view of African genesis of abelisaurids and titanosaurids is supported by another line of evidence: the vector of dispersal between continents generally takes place from larger (o smaller areas. Faunal interchange occurred between Europe and Africa much of the Cretaceous time across the Tethys, hetween northwestern Africa and Theria, possibly via the Alboran plate, or the Apulian plate”, The abelisawid — Turacusuuruy and titanosaurids Ampelosaurus and Magyarosaurus in Europe certainly attests to intermittent connections between Europe and Africa during that time by island hopping Jurassic form One can Death of the Dinosaurs Ruling the Earth for 167 million years, dinosaurs seemed indestructible. Yet. they disappeared suddenly at the end of the Cretaceous, some 65 million years ago. The KT extinction brought to an SANKAR CHATTERJEE AND CHRISTOPHER R SCOTESE end not only dinosaurs, but other life forms as well, incnding plants and two-thirds af marine species. What killed the dinosaurs? Currently, two catastrophic models have been proposed to explain, the demise of the dinosaurs: the volcanic hypothesis and the meteorite impact hypothesis. The volcanic hypothesis links extinction to massive Deccan volcanism in India that occurred around the KT boundary“. Could the pollutants from the Deccan eruption have damaged the environment and triggered the extinction? This is certainly possible, albeit indirectly, ‘he large areal extent of the lava flows certainly affected the local flora and fauna by habitat destruction and pollution. However, field evidence does not support any extinction event of local biota in the trappean beds between the lava tlows*. It appears that Deccan volcanism had little direct effect on the demise of the dinosaurs on a global scale. The more direct cause appears to be the impact of a large meteorite, which struck the Earth and wiped out dinosaurs and other animals In 1980, the Alvarez group discovered a thin layer of iridium-rich clay in many parts of world which coincided with the extinction event of the dinosaurs. Iridium is rare on the Earth's surface, but it is often found in a chondritic meteorite. The Alvarez. group proposed that the iridium layer was the result of the impact of a giant meteorite, about 10 km in diameter. The recent discovery of the actual point of collision of this killer meteorite, the Chicxulub crater in the Yucatan Peninsula of Mexico. has bolstered the impact hypothesis’. The impact carved out a tremendous crater, about 180 kun int diameter, which is now buried under a thick pile of sediment. Recently, a near-synchronous, almost antipodal, even larger impact crater named Shiva, has been identified off the west coast of India‘. Centered on a structural uplift called the Bombay High, a giant oil basin on the continental shelf of western India, the 600 km-long, 450 km-wide Shiva crater was presumably split into two parts by the Carlsberg Ridge which separated India from the Seychelles Island 65 million years ago. One part of the crater is presently attached to the Seychelles and the other half to the western India. The evidence of the Shiva impact event seems to accord well with age and morphology of the crater. The oblong shape of the crater and the asymmetric disuibution uf MuidGONDWANA. EVOLUTION AND BIOGEOGRAPHY OF THE INDIAN PLATE ejecta indicate oblique impact in a SW-NE trajectory. It is estimated that a 40-km asteroid crashed on the western coast of India around 65 Ma, excavating the Shiva crater, shattering the lithosphere, and triggered the India-Seychelles rifting, How devastating was the collision and how did it affect the life? Various models have been proposed in recent times to explain the killing mechanism™ The impact generated a force thousands of times more powerful than the explosion of all nuclear arsenal in the world, It injected a large volume of dust into atmosphere that blocked sunlight for months and prevented photosynthesis; as a result, plants died, followed by herbivores, and eventually the food chain collapsed, and all dinosaurs perished. The tremendous impact would have ignited global fires, initiated tsunamis, destroyed coastal habitats, produced acid rains, tumed seawater acidic, dissolved carbonate-shelled animals, and devastated biosphere. Millions of organisms would have died instantly from the tremendous tremor and the global fire generated from the impact. It was a holocaust of unmatched dimensions in the geologic history. Are the dinosaurs really extinct? Ina phylogenctic sense, probably not. Currently, birds are considered to be living dinosaurs, the direct descendants of theropods”. If so, dinosaurs did not die out completely. While it is true that all tenestiial form ae wiped out completely at the KT boundary, one group of dinosaurs escaped this catastrophe. They are still around us-~flying, hopping, perching, singing, and nesting. They inspired us to fly and invent the airplane. We call them birds. After the dinosaur extinction, mammals underwent an explosive evolution in the beginning of the Tertiary, diversifying and adapting to many different ecological niches. As a result, the distribution mammalian fossils provide important biogeographic tool to check the paleocontinental positions. It helps to assess the timing of India~ Asia collision. India-Asia Col ion Age: Paleontologic Evidence There is no fossil record of terrestrial Paleocene vertebrates from India. The distribution of marine 423 Ranikot Formation and its equivalents at the northern edge of the drifting Indian plate indicates that it was surrounded by sea. The plate tectonic reconstruction map (60 Ma, Fig. 14) supports this contention. During this time, India was separated from Asia hy a wide stretch of Tethys Sea. However, it may have maintained its intermittent contact with Africa through the Greater Somalia. During the Middle Eocene (50 Ma, Fig. 15), India established a narrow filter bridge at its Northeastern margin with Asia. This dispersal route allowed for the first time the great influx of Asian mammals to India. This is the initial stage of the collision of India with Asia, corroborated by both palaeontological” and geophysical evidence”. The fossil record indicates that there was no biologic isolation of India from Asia during this period. The oldest large mammals recorded from the Indian subcontinent are from Mid Eocene of Pakistan, Kutch, Jammu and Kashmir. In Pakistan, the Ganda Kas and Lamindhan localities have a mammalian fauna of taeniodonts, creodonts, titanotheres, anthracotheres, artiodactyls, and perissodactyls from the Charat Series; all are very similar to Mongolian, Burmese and and North American taxa”. In the Sabathu Formation of Jammu and Kashmir, Sahni and Khare” recorded a tapiroid, bunodont artiodactyls, a paramyid rodent, and a hyracodont rhinoceratid, all closely related to Asian and North American forms. Krom Kutch, Sahni and Misra” reported a cetacean Protocetus, also known from Egypt and North America, and a primitive mesonychid camivore Harpogolostes, recorded also from Pakistan, Mongolia, and North America. Other evidence for Asian contact is indicated by primates, brontotheres, and pristichampsid crocodiles. However, anthracobunid proboscideans and —dyrosaurid crocodiles show affinities with those of African continent”, probably through the Greater Somalia. Conclusions The physical and biotic links of India during the breakup of Gondwana and its subsequent union with Asia is one of the greatest puzzles in plate tectonics and hiogeography This puzzle can he solved using two lines of evidence. The tectonic evolution of the Indian plate from its Pangean origin to the present day based on geological and geophysical evidence (Figs.1-14). Though isolated