GOLDEN JUBILEE MEMOIR OF THE GEOLOGICAL SOCIETY OF INDIA

No.66, 2008, pp.203-225

Geodynamic Evolution of the Indian Plate:

Consequences for Dispersal and
Distribution of Biota

ASHOK SAHNI1 and G. V. R. PRASAD2

1
CAS in Geology, Panjab University, Chandigarh – 160 014,
2
Department of Geology, University of Jammu, Jammu – 180 006,
Email: ashok.sahni@gmail.com; guntupalli.vrprasad@gmail.com

Abstract
The Indian plate underwent a prolonged period of physical isolation from
the time of its break-up from the former Gondwanaland until its collision with
Asia. India’s northward journey involved four phases: rifting and early phase of
drifting, late phase of drifting, pre-collision phase and collision phase. Fossil
data from the early phase of drifting is meager because of very few
palaeontological studies on Early Cretaceous continental deposits. Extensive
research in recent years on the Upper Cretaceous Deccan infra-and intertrappean
beds and Early Eocene deposits of India led to the recovery of diverse groups of
vertebrates. Contrary to the evolution of endemic fauna and flora during its
northward flight, fossil evidence from different phases of drifting offers a complex
biogeographic picture. The fossil data from the Late Cretaceous demonstrates
biogeographic affinities with both Gondwanan and Laurasian landmasses besides
endemic evolution of some taxa, whereas the biota from the Early Eocene is
represented by both endemic and Holarctic taxa. Further, Late Cretaceous
vertebrates favour the continuance of a terrestrial link between South America
and India-Madagascar as late as 65 Ma ago. The anomalous occurrence of
Laurasian biotic elements in the Late Cretaceous of India is explained by dispersals
across island-arc systems by sweepstakes mode of dispersal. The presence of
endemic taxa in addition to Holarctic fauna in the Early and Middle Eocene
times is suggestive of the fact that the Indian subcontinent was partially isolated
by an epicontinental sea permitting only limited faunal interchanges with Asia.
There is also limited fossil evidence for Out-of-India dispersal hypothesis. Recent
finds of oldest occurrences of grasses, freshwater diatoms, oldest ungulate-like
mammal, whales, and archontans favour India as a centre of origin for many
animal and plant groups.
204 ASHOK SAHNI AND G. V. R. PRASAD

INTRODUCTION

The geographic distribution of organisms and global biodiversity patterns

depend on factors like climate, latitudinal position and geodynamic history of
the landmasses. Consequent upon the break-up of former Gondwanaland,
landmasses which were previously continuous or contiguous became disjunct,
and new bodies of marine water were created. This redistribution of landmasses
and seaways (rifting, drifting, and recombination) greatly affected the free
movement of terrestrial and marine organisms because of the changed climatic
and environmental conditions. Though the Indian plate is the best example
one can cite for such tectonic events, the effects of these events on terrestrial
vertebrate evolution in the Indian subcontinent remained least understood. Plate
tectonics envisaged a long period of isolation for the Indian plate from its Late
Jurassic separation from Africa to its collision with Asia in the early Tertiary.
Traditionally, palaeogeographic maps depicted the following sequence of plate
tectonic events for the dismemberment of the former Gondwanaland (Powell,
1979; Smith et al. 1994; Storey et al. 1995; Plummer, 1995; Subramanya, 1998,
Hay et al. 1999; Pande et al. 2001).

30 Ma - Break-up of South America and Antarctica

55-53 Ma - Collision of India with Asia
65 Ma - Rifting of India from the Seychelles contemporaneous with the eruption
of Deccan flood basalts of India
90-85 Ma - Break-up of Madagascar from India and the Seychelles. This event
coincided with the eruption of flood basalts and associated rhyodacites
from the eastern margin of Madagascar, igneous rocks from the Seychelles
microcontinent, St. Mary’s Islands volcanics
95 Ma - Rifting of Australia, Antarctica and New Zealand
100 Ma - Separation of South America and Africa
130-120 Ma - Separation of Australia – Antarctica from India-Madagascar, and the
Seychelles more or less coinciding with Kerguelen basalts (118 Ma) and
Rajmahal basalts (115-118 Ma) of India.
160-150 Ma - Break-up of East Gondwanaland (Australia, Antarctica, Madagascar,
India and the Seychelles) from eastern part of West Gondwanaland
(eastern Africa).

Plate tectonics, therefore, favour a long period of isolation for the Indian
subcontinent, which is sometimes estimated at 100 Ma (Briggs, 1989). Even if
we consider the time of break-up with Madagascar at about 88 Ma as the
beginning of India’s isolation, the Indian plate would have underwent physical
isolation for about 30 Ma until its collision with Asia around 55 Ma. If it
experienced a prolonged period of isolation from other landmasses, one would
expect to find evidence for the evolution of endemic fauna and flora on the
Indian subcontinent during its northward flight. Now the question is where
 GEODYNAMIC EVOLUTION OF THE INDIAN PLATE 205

should we look for such evidence? As expected the fossil vertebrates from the
continental Triassic and Jurassic strata of India exhibit cosmopolitan distribution
(Jain and Bandyopadhyay, 1997) because the Gondwanan and Laurasian
continents were still in close contact. Therefore, for testing the endemicity of
biota during the northward flight of India, the Cretaceous and early Tertiary
biota is crucial as it was during the Cretaceous-Early Eocene period that rapid
northward drift of the Indian plate took place. As they cannot negotiate wide
bodies of salt water, the continental fauna and flora would play a significant
role in understanding the contiguity or distantness of landmasses. Hence the
continental Cretaceous - Eocene fossil record of India is very important for
understanding the effects of plate tectonics on the biota. In light of recent
researches on continental Cretaceous – Eocene faunas from the Indian
subcontinent, we present here an overview of India’s palaeobiogeography during
its northward flight.

BIOTIC DISPERSALS IN A PLATE TECTONIC SETTING

Before we discuss various palaeobiogeographic models, a brief
introduction to different modes of faunal dispersals is necessary. In a classic
paper, McKenna (1973) discussed the consequences of fragmentation and
recombination of landmasses on pre-existing biotas at a great length. He
visualized three modes of biotic dispersals in a plate tectonic setting viz.,
corridor, filter bridges, and sweepstakes dispersals. According to McKenna
(1973), a corridor serves as a route for the spread of many or most of the
animals between two connected areas. The best example for this is the Central
Asian terrestrial corridor linking the faunas of Europe and China since 30 Ma.
Filter bridges form a series of lesser barriers within a limited variety of habitats
causing differential filtering of certain organisms as against others. In such a
setting dispersals take place both ways and are not totally unbalanced. The
tropical lowlands of isthmus of Panama are one example for the filter route.
Sweepstakes, the third mode of dispersal, is random, piecemeal, unpredictable,
and takes place over long distance of water. The fauna may be unbalanced and
dispersals may be sporadic as across the oceanic barriers between Pacific islands.
As visualized by McKenna (1973), when a landmass splits apart, the mode of
biotic dispersal changes from an initial continuous terrestrial corridor to a filter
bridge and finally to a phase of sweepstakes. This sequence is reversed when
two landmasses come together.

NORTHWARD FLIGHT OF INDIA: FOSSIL EVIDENCE

The northward journey of the Indian plate involved four important
206 ASHOK SAHNI AND G. V. R. PRASAD

phases: (1) Rifting and Early Phase of Drifting, (2) Late Phase of Drifting,
(3) Pre-Collision Phase, and (4) Collision/Post-Collision Phase.

Rifting and Early Phase of Drifting (160-80 Ma)

When rifting is initiated between adjacent landmasses, multiple ecosystems
are created in the near shore environments leading to diversification of estuarine
and marine organisms. With the onset of drifting, commonality of species
between once adjacent landmasses decreases. Geophysical data favoured a
drift rate of 3-5 cm/year for the Indian plate between 130 Ma to 80 Ma (Powell,
1979). The fossil record from the rifting and early drifting phases (160-80 Ma)
of India is primarily known from the continental Jurassic of India which exhibits
both Laurasian and Gondwanan affinities (Jain and Bandyopadhyay, 1997;
Prasad and Manhas, 2002). However, there is also some evidence for supposed
endemic evolution during the Jurassic period. DNA sequencing of
Nasikabatrachus sahyadrensis (Nasikabatrachidae), a new frog reported from
the Western Ghats region (Biju and Bossuyt, 2003; Dutta et al. 2004), indicated
that it is an endemic taxon that split off from its sister taxon Sooglossidae of
the Seychelles island in the Jurassic (150 Ma). As the divergence of
Nasikabatrachidae of India from Sooglossidae of the Seychelles took place
much before the break-up of India-Seychelles block at about 65 Ma, it was
hypothesized that shrinkage of table lands following the rise in Jurassic sea
level may have caused isolation of these two sister taxa (Dutta et al. 2004).
On the other hand, the fossil record from the Early Cretaceous and early
Late Cretaceous of India is very limited. The Lower Cretaceous Dhrangdhara
and Athgarh formations and east coast Gondwana deposits of peninsular India
are yet to yield vertebrate fossils. The microvertebrate fauna of Lower
Cretaceous Gangapur Formation is represented by ganoid fishes of little
palaeobiogeographic significance (Prasad et al. 2004). Petrified tree trunks,
angiosperms of mangrove ecosystem (Rhizophora) and sauropod dinosaur
remains with close affinities to Saltasaurus of Africa have been documented
from the Nimar Formation which underlies the marine Bagh Formation of
Turonian-Coniacian age (Khosla et al. 2003). The fauna from the Bagh
Formation is mainly represented by marine organisms. The ammonite fauna of
Bagh Formation compares very well with that of Zululand and Madagascar
implying that a single marine biogeographic province existed between these
landmasses until the Coniacian (Bardhan et al. 2002).

Late Phase of Drifting (Late Cretaceous, 80-65 Ma)

During the late phase of drifting, the Indian plate underwent rapid
northward flight at a rate of 15-20 cm/year (Powell, 1979). When landmasses
undergo complete physical isolation, increased endemism of the constituent
 GEODYNAMIC EVOLUTION OF THE INDIAN PLATE 207

populations, evolution and diversification of new taxa is expected as it is evident

in case of Madagascar, Australia, and South America. Does the fossil record
from the late phase of India’s northward flight reflect this? By Maastrichtian,
the marine ammonite fauna of India became highly endemic even at species
level (Bardhan et al. 2002) pointing to complete physical isolation of India
from other Gondwanan landmasses. As compared to the fossil record of early
phase of drifting, it is better documented from the late phase of drifting
represented by the biota from the Upper Cretaceous (Maastrichtian) Kallamedu
Formation and the Upper Cretaceous (Maastrichtian) Deccan infra- and
intertrappean beds.
The Kallamedu Formation of Cauvery basin in South India represents a
continental facies consisting of sandstones, clays and sandy clays and occurs
sandwiched between shallow marine Globotruncana gansseri – bearing
Kallankurichchi Formation and Early Palaeocene Niniyur Formation (Tripathi
and Mamgain, 1987) (Fig.1). The Kallamedu Formation is long known for its
dinosaur yielding bone beds. Unfortunately, most of the reported skeletal
remains from this formation are fragmentary in nature and their assignment to
titanosaurids, megalosaurids, and stegosaurids is debatable (Chatterjee and
Rudra, 1996). Gaffney et al. (2001) described a side-necked turtle, Kurmademys
kallamedensis belonging to the family Bothremydidae (Pelomedusoides), a
group known essentially from the former Gondwanaland, from the Kallamedu
Formation.
In the last two decades, application of bulk screen-washing techniques to
the Deccan infra- and intertrappean beds (Fig.1) has significantly improved
the Late Cretaceous continental vertebrate fossil record (Khosla and Sahni,
2003 and references therein). The vertebrate fauna recovered from these
beds includes fishes, frogs, lizards, snakes, turtles, crocodiles, dinosaurs, and
mammals. The Deccan infra- and intertrappean fishes (Lepisosteus,
Osteoglossiformes, Siluriformes, Pycnodontiformes) are not taken into account
for palaeobiogeographic reconstructions because of their cosmopolitan
distribution. As compared to this, pelobatid (Sahni et al. 1982), discoglossid,
hylid (Prasad and Rage, 1991, 1995), Myobatrachinae (Noble, 1930; Spinar
and Hodrova, 1985), leptodactylid and ranoid frogs (Prasad and Rage, 2004);
anguimorph lizards (Prasad and Rage, 1995); nigerophiid (Rage and Prasad,
1992) and madtsoiid (Rage et al. 2004) snakes; pelomedusoid turtles (Jain,
1977, 1986; Bajpai et al. 1990; Gaffney et al. 2003); mesosuchid crocodiles
(Prasad and De Broin De Lapparent, 2002); abelisaurid dinosaurs (Chatterjee,
1978; Wilson et al. 2003); and gondwanathere (Das Sarma et al. 1995; Krause
et al. 1997; Prasad et al. 2007a, Wilson et al. 2007), haramiyid (Anantharaman
et al. 2006), and eutherian (Prasad and Sahni, 1988; Prasad et al.1994; Prasad
and Godinot, 1994; Godinot and Prasad, 1994; Rana and Wilson, 2003; Khosla
208 ASHOK SAHNI AND G. V. R. PRASAD

Fig.1. Map of the Indian subcontinent showing Cretaceous and Eocene vertebrate yielding
sites.

et al. 2004; Prasad et al. 2007b) mammals figured prominently in most of the
palaeobiogeographic reconstructions (Sahni, 1984; Sahni et al.1987; Jaeger
et al. 1989; Briggs, 1989, 2003; Sahni and Bajpai, 1991; Prasad and Rage,
1991; Chatterjee, 1992; Prasad et al.1995; Krause et al. 1997; Chatterjee and
Scotese,1999; Sahni and Prasad, 1999; Prasad and Sahni, 1999; Rage, 2003;
Prasad, 2005; Sahni, 2006). Contrary to endemic fauna expected from the
northward drifting Indian plate, the fauna exhibits, close biogeographic affinities
with both Laurasian and Gondwanan taxa. As the taxonomic affinities of these
vertebrate groups has been discussed in great detail in the above cited works,
only a brief summary of their biogeographic affinities is given here. Among
the Deccan infra- and intertrappean vertebrate taxa, myobatrachid, leptodactylid,
 GEODYNAMIC EVOLUTION OF THE INDIAN PLATE 209

hylid, and ranoid frogs, pelomedusoid turtles, mesosuchid crocodiles,

nigerophiid snakes, abelisaurid dinosaurs, haramiyid and gondwanathere
mammals have typical Gondwanan distribution. The recent report of a
baurusuchid crocodile of South American affinity from the Upper Cretaceous
Pab Formation of Baluchistan (Pakistan) is another addition to this list of
Gondwanan taxa (Wilson et al. 2001). On the other hand, pelobatid and
discoglossid frogs, anguimorph lizards and eutherian mammals exhibit close
biogeographic affinities to those of Eurasia. Additional evidence for Laurasian
biogeographic relationship comes from charophytes (Bhatia et al.1990,
Srinivasan et al. 1994).
In addition to the vertebrate fauna, freshwater ostracods have also been
reported from the Deccan infra- and intertrappean beds. All the early works
suggested Eurasian affinities for the ostracod fauna (Bhatia and Rana, 1984;
Bhatia et al. 1996; Sahni and Khosla, 1994; Bhandari and Colin, 1999, Khosla
and Sahni, 2000). In a series of papers published in the last five years, as many
as 100 species of ostracods have been described from the intertrappean beds of
Kachchh, Nagpur, Gurmatkal, Mohagaonkalan and the taxonomic status of
many species described in earlier works has been revised (Whatley and Bajpai,
2006 and references therein). Though at generic level many of these taxa (about
39%) exhibited Mongolian and Chinese affinities, at species level they were
highly endemic (Whatley and Bajpai, 2006). However, two species are also
known from the Upper Cretaceous strata of Mongolia and China. Cypridea
cavernosa was earlier recorded from Russia, China and Mongolia, while
Frambocythere tumiensis is known by a different subspecies in Africa,
China, France and Spain.

Pre-Collision Phase (Early Eocene, 55-53 Ma)

In the northern landmasses, the Early Eocene, one of the warmest periods
of the Cenozoic Era, has witnessed rapid diversification of land mammals. In
the Indian subcontinent, Neotethys underwent shallowing and extensive coal
swamps developed in the western region extending from Baluchistan in
Pakistan (Ghazij Formation) to Vastan Lignite Mine in Surat District, Gujarat,
and Giral Lignite Mine (Akli Formation), Barmer District and Palana Lignite
of Bikaner District, Rjasthan (Fig.1). Amongst these, only Ghazij Formation
and Cambay Shale of Vastan Lignite Mine have yielded highly diversified
mammalian assemblages that offer an insight into the vertebrate groups present
on the Indian raft. At least 11 orders of placental mammals and two taxa
belonging to marsupials in addition to fishes, amphibians, lizards, snakes,
crocodiles, turtles and birds, representing the oldest Cenozoic vertebrate fauna
of South Asia have been documented so far from the Vastan Lignite Mine. The
placental orders documented from this Early Eocene site include Proteutheria,
210 ASHOK SAHNI AND G. V. R. PRASAD

Apatotheria, Insectivora, Artiodactyla, Perissodactyla, Primates, Rodentia,

Lagomorpha, Chiroptera, Creodonta, and Arctocyonia, many of which have
been reported for the first time from India (Bajpai et al. 2005a, b, c, 2006,
2007; Rana et al. 2005; Rose et al. 2006; Smith et al. 2007; Rana et al. 2008;
Rose et al. 2008). This fauna is an admixture of endemic and Holarctic taxa.
Vastanavis eocaena, the oldest fossil bird from the Indian subcontinent (Mayr
et al. 2007) is distinct from known fossil birds from the Eocene of Europe and
North America and appears to be endemic to India. The artiodactyls (Gujaratia
indica, G. pakistensis), perissodactyls (Cambaytherium), and primates
(Marcgodinotius, Vastanomys) and anthracobunids (Indobune) also represent
the endemic forms, whereas insectivores (Vastania, Cambaya), apatotherians
(Frugivastodon), cimolestids (Suratilestes) and palaeoryctids (Anthraryctes)
have closely related forms from the Holarctic continents (Bajpai et al. 2005a).
Recently, a highly diversified bat fauna represented by seven genera and seven
species with strong affinities with European bat faunas of Paris basin (France)
and Messel locality of Germany has been reported from Vastan Lignite mine
(Smith et al. 2007). In marked contrast, the mammal fauna from the lower and
middle parts of Ghazij Formation is endemic, while those from the upper part
of this formation are predominantly Holarctic and may have reached the Indian
subcontinent via a series intermittently emergent islands (Gingerich et al. 1997;
Clyde et al. 2003).

Collision/Post-Collision Phase (Middle Eocene, 49 Ma)

The subaerial contact between the Indian plate and Asian mainland was
marked by complete withdrawl of epicontinental sea (Neotethys) from the
northern margin and its restriction to the western and eastern margins, reversal
of drainage as indicated by change in palaeocurrent directions, climatic and
ecosystem changes that followed the rise of Himalaya and concomitant
increase in clastic inputs (Sahni, 2006). The closure of Tethys was followed by
depletion of shelf ecospace severely affecting the marine biodiversity and
establishment of terrestrial corridor between the two colliding landmasses. With
the creation of stable corridor, many Asiatic elements like primitive rhinoceros,
tapiroids, brontotheres, and ctenodactylid rodents could disperse from Asian
mainland. The vertebrate fauna from the early Middle Eocene Kuldana
Formation of Pakistan and Subathu Formation of India (Fig.1) provide us with
insights into the diversity and palaeobiogeographic relationships of the fauna
present during the collision phase. The Middle Eocene ecosystems of India
harboured many endemic mammalian groups such as tethytheres,
anthracobunids, raoellid artiodactyls, and perissodactyls. Origin and evolution
of whales, their transition from terrestrial to marine aquatic life also took
place on the Indian subcontinent during this phase. This is one of the best
 GEODYNAMIC EVOLUTION OF THE INDIAN PLATE 211

examples of macroevolution. Many Eurasian taxa like ctenodactylid rodents,

represented by Chappatimyidae, a family having a subcontinental distribution
(Kumar et al. 1997) also made their appearance possibly utilizing the newly
established terrestrial connection. When compared to later Murree fauna
which was dominated by North American, European, and Asiatic forms, the
Subathu fauna is more endemic in nature (Kumar and Kad, 2003). The
development of some endemic taxa in the Kuldana and Subathu formations
points to the fact that at the time of their deposition the Indian subcontinent
was partially isolated from adjoining areas by an epicontinental sea allowing
limited faunal interchanges (Thewissen et al. 2001).

PALAEOBIOGEOGRAPHIC MODELS
As enumerated above, the Indian plate hosted both endemic and non-
endemic (Laurasian or Gondwanan) taxa during different phases of its journey
towards the north. However, the Late Cretaceous phase (Late Phase of Drifting)
has attracted more attention than any other phase because of extensive research
carried out on the Deccan infra- and intertrappean biota in recent years and the
presence of both Laurasiatic and Gondwanan taxa in addition to some endemic
taxa. To explain this palaeobiogeographic anomaly, a number of working
hypotheses have been proposed. These include: (a) Gondwanan Connection
and (b) Laurasian Connection (Dispersals via NE Africa, Early India/Asia
collision, Sweepstakes dispersals). In the following section, we discuss these
hypotheses in some detail.

Gondwanan Connection
Sahni (1984) and Sahni et al. (1987) were the first to throw light on the
Gondwanan affinities of Late Cretaceous biota of India. As the prevailing
geophysical data favoured the break-up between Antarctica-Australia and
India-Madagascar-Seychelles at 130-120 Ma (Powell, 1979; Patriat and
Achache, 1984), they argued that aseismic structures like Mascarene Plateau
and Chagos-Laccadive ridges may have served as intervening terrestrial
link between India, Madagascar and South America. Strong evidence for a
terrestrial link between Indo-Madagascar and South America comes from
vertebrate fossils such as abelisaurid dinosaurs, baurusuchid crocodiles, and
gondwanathere mammals (Krause et al.1997; Wilson et al. 2001, 2003; Prasad
et al. 2007a; Wilson et al. 2007). Rajasaurus narmadensis Wilson et al., 2003,
an abelisaurid dinosaur reported from the Lameta Formation was regarded as a
sister taxon of Majungatholus from the Upper Cretaceous Maevarano
Formation of Madagascar and Carnotaurus from the Upper Cretaceous of
Argentina. Similarly, baurusuchid crocodile Pabwehshi pakistanensis Wilson
212 ASHOK SAHNI AND G. V. R. PRASAD

et al. 2001, documented from the Upper Cretaceous Pab Formation of

Baluchistan (Pakistan), represents the only baurusuchid crocodile known outside
South America. Gondwanatherian mammals represent an enigmatic group of
mammals with high-crowned teeth known from the former Gondwanaland
(Bonaparte, 1990; Krause et al.1997; Krause et al. 2003; Goin et al. 2006;
Prasad et al. 2007a; Wilson et al. 2007). The Indian gondwanathere species
Bharattherium bonapartei Prasad et al., 2007a (Fig.2a-j) is a sister taxon of
Lavanify miolaka Krause et al. 1997 reported from the Upper Cretaceous
Maevarano Formation of Madagascar and is closely related to Campanian
Sudmamerica of Argentina. The recent report of a gondwanathere mammal in
the Cretaceous sediments of Tanzania implies that either gondwanatheres
were widely distributed in the former Gondwanaland prior to the break-up of
South America and Africa or terrestrial links were maintained between Africa

Fig.2a-j. Cheek teeth of Bharattherium bonapartei Prasad et al. 2007a. a-d, holotype (VPL/
JU/IM/33) in lateral (a-b), enlarged lateral view of b showing growth lines (c), and
occlusal views (d) (Upper Cretaceous intertrappean beds of Kisalpuri). e-j, referred
specimen (VPL/JU/NKIM/25) in lateral (e-h), occlusal (i), and enlarged occlusal views
(j) (Upper Cretaceous intertrappean beds of Naskal) (Prasad et al. 2007).
 GEODYNAMIC EVOLUTION OF THE INDIAN PLATE 213

and other Gondwanan continents well into Late Cretaceous. If an Upper

Cretaceous age is confirmed for the gondwanathere yielding strata of Tanzania
by further studies, it will negate Africa first hypothesis (Sampson et al.1998)
which assumes that Africa remained isolated from other Gondwanan continents
after its separation from South America in the Early Cretaceous. In addition to
these three vertebrate groups, Indobatrachus (Myobatrachinae), leptodactylid
(Fig.3a-c), hylid, and ranoid frogs, nigerophiid (Fig.4a-e) and madtsoiid snakes
(Fig.5a-e), and pelomedusoid turtles also support the Gondwanan connection.
These latter groups and gondwanathere mammals being small in size may not

Fig.3a-c. Ilium (VPL/JU/1021) of Leptdactylidae frog from the Upper Cretaceous intertrappean
beds of Naskal in lateral (a), anterior (b), and posterior (c) views (after Rage and Prasad,
2004).

Fig.4a-e. Mid trunk vertebra (VPL/JU/500) of Indophis sahnii Rage & Prasad, 1992 from the
Upper Cretaceous intertrappean beds of Naskal in ventral (a), left lateral (b), dorsal (c),
anterior (d), and posterior (e) views (after Rage and Prasad, 1992).
214 ASHOK SAHNI AND G. V. R. PRASAD

Fig.5a-e. Posterior trunk vertebra (VPL/JU/1519) of Madtsoiidae snake from the Upper Cretacous
intertrappean beds of Kelapur in dorsal (a), left lateral (b), ventral (c), anterior (d), and
posterior (e) views.

have needed a continuous terrestrial route. But the dispersal of large abelisaurid
dinosaurs and baurusuchid crocodiles would have definitely required a land
corridor or a filter bridge. In conformity with the fossil evidence (Krause et
al.1997), the recent palaeogeographic data attested to the presence of a terrestrial
link between Indo-Madagascar-Seychelles block and South America via
Kerguelen Plateau (Storey et al. 1995; Hay et al. 1999) as late as 80 Ma ago.
Case (2002), on the other hand, favoured a similar terrestrial connection across
Gunnerus Ridge instead of Kerguelen Plateau..

Laurasian Connection
Different biogeographic models have been proposed to account for the
presence of Laurasian taxa, such as pelobatid and discoglossid (Gobiatinae)
frogs (Fig.6a-d), anguimorph lizards, and eutherian mammals (Fig.7a-f) in the
Late Cretaceous India. These include: (1) dispersals from northeast Africa of
Briggs (1989, 2003), and Chatterjee (1992) and Chatterjee and Scotese (1999),
(2) early India/Asia collision of Jaeger et al. (1989), and (3) sweepstakes
dispersals of Prasad and Sahni (1999).

Dispersals via NE Africa: Based on the lack of endemism in the Late

Cretaceous vertebrates of India and presence of dinosaurs and other vertebrates
of both Gondwanan and Laurasian affinities, Briggs (1989, 2003) suggested
that in the Early Cretaceous, the Indian plate was isolated but by Middle
Cretaceous it came close to NE Africa filling the gap between Somalia and
Eurasia and permitting biotic dispersals from Africa as well as Eurasia. His
conclusions were primarily based on the biogeographic affinities of plant,
invertebrate and vertebrate fossils from the Jurassic Kota Formation and
 GEODYNAMIC EVOLUTION OF THE INDIAN PLATE 215

Fig.6a-d. Ilia of Gobiatinae (Discoglossidae) frogs from the Upper Cretaceous intertrappean
beds of Naskal. a. right ilium (VPL/JU/1016) in lateral view, b.left ilium (VPL/JU/
1017) in lateral view, and c-d. right ilium (VPL/JU/1018) in lateral (c ) and anterior (d)
views (after Rage et al. 2004).

Cretaceous Deccan intertrappean beds. A slightly modified biogeographic model

was proposed by Chatterjee (1992) and Chatterjee and Scotese (1999).
According to this model, Somalia and India were closely placed in the Late
Cretaceous allowing dispersal of continental vertebrates between Africa and
India and a northern connection existed through Dasht-i-Marg (Central

Fig.7a-f. Upper teeth of Deccanolestes hilopi Prasad & Sahni, 1988. a-b. holotype, right M1
(VPL/JU/NKIM/10) in occlusal (a) and anterior (b) views. c-d. right P3 (VPL/JU/NKIM/
19) in lingual (c) and occlusal (d) views. e-f. right M3 (VPL/JU/NKIM/11) in occlusal
(e) and anterior (f) views (after Prasad et al. 1994).
216 ASHOK SAHNI AND G. V. R. PRASAD

Afghanistan) and Lut block (east-central Iran) and Kashmir region (India)
facilitating dispersals from Eurasia. The NE African connection appears very
attractive in light of recent discovery of a gondwanathere mammal in the
Cretaceous of Tanzania (Krause et al. 2003) and this may explain the late
survival of similar Gondwanan taxa both in India and Laurasia. However, this
needs to be authenticated through the recovery of common terrestrial faunal
elements from Africa and India. Unfortunately, the fossil record from the Upper
Cretaceous strata of Africa is impoverished, whereas Early Cretaceous
record is well documented. In comparison to this, the Upper Cretaceous fossil
record of South America, India, and Madagascar is the best documented among
the Gondwanan continents, while the Early Cretaceous fauna from these
landmasses is hardly known. Therefore, to test the NE African terrestrial
connection, the fossil record from the Upper Cretaceous rocks of Africa
needs to be improved.

Early India/Asia Collision: Contrary to widely accepted Middle Eocene

age for India/Asia collision, Jaeger et al. (1989) suggested that subaerial contact
between Greater India and the Asian plate was established around Cretaceous-
Tertiary boundary. This was based on Late Cretaceous - Early Palaeocene
deformation history, crustal shortening and thickening in Asia and the presence
of pelobatid frogs and eutherian mammal Deccanolestes of Laurasian affinity
in the Late Cretaceous of India. Geophysical data from the Indian Ocean and
Chitral region did provide some support to this hypothesis (Klootwijk et al.1992,
1994). However, stratigraphic, sedimentological and tectonic evidences from
the northern passive margin of India do not support the early India/Asia collision
hypothesis. Marine sedimentation continued along the northern margin of India
well into Early Eocene (Gaetani and Garzanti, 1991; Garzanti et al. 1987; Searle
et al. 1997). Moreover, a vast majority of dates for India/Asia collision cluster
around 53-55 Ma (Powell, 1979; Patriat and Achache, 1984; Besse and
Courtillot, 1988; Gaetani and Garzanti, 1991; Garzanti et al. 1987; Searle et al.
1997) and even younger dates of 51.8 - 45 Ma (Rowley, 1997) and 34 Ma
(Aitchison et al. 2007) were suggested recently. A slightly modified version of
early India/Asia collision was offered by Rage (1996, 2003). On the basis of
past and present distribution of boine snakes and iguanid lizards, Rage (2003)
suggested a terrestrial route between Asia and Madagascar through India and
the Seychelles that enabled dispersal of boine snakes, iguanid lizards and
lemuriform primates

Sweepstakes Dispersals: As there is no convincing evidence to show

that India/Asia collision took place close to the Cretaceous-Tertiary boundary,
sweepstakes dispersals have been invoked to account for the presence of
 GEODYNAMIC EVOLUTION OF THE INDIAN PLATE 217

Laurasian taxa like pelobatid and discoglossid frogs, anguimorph lizards,

eutherian mammals, and charophytes in the Late Cretaceous of India at a time
when India was still separated from Asia by a considerable body of marine
water (Sahni and Bajpai, 1991; Prasad and Sahni, 1999; Prasad, 2005). In
comparison to Gondwanan abelisaurid dinosaurs and baurusuchid crocodiles
which needed stable terrestrial corridor for dispersal, the Late Cretaceous
Laurasian taxa of India are very small in size approaching the size of a house
rat. Such a small size could have been advantageous for rafting or waif
dispersal. Prasad and Sahni (1999) argued that Kohistan-Dras island-arcs and
Trans-Himalayan magmatic arc which occupied intermediate position between
the Indian plate and Asia and several other unknown oceanic islands which
did not leave any physical evidence may have facilitated sweepstakes dispersals
or island hopping between India and Asia. This is the mode of dispersal one
would expect between landmasses that are about to collide (McKenna, 1973).

OUT-OF-INDIA DISPERSALS
In a classic paper, Krause and Maas (1990) expounded a new idea that
ancestral stocks of perissodactyls, artiodactyls, and primates arrived in India
via Madagascar, evolved in isolation on the Indian subcontinent during its
northward journey, and finally dispersed out of India into Asia when subaerial
contact was established between these two landmasses. This is more or less
similar to what is known today as Out-of-India dispersal hypothesis (Bossuyt
and Milinkovitch, 2001). The latter hypothesis assumes that certain extant
Asian taxa having ancient Gondwanan origin reached Asia by rafting the
northward drifting Indian plate. The Out-of-India dispersal hypothesis was also
suggested for aplocheiloid (Murphy and Collier, 1997), Asian arowana
(Kumazawa and Nishida, 2000) and cichlid (Sparks, 2004) fishes; acrodont
lizards (Macey et al. 2000); ranoid frogs (Bossuyt and Milinkovitch, 2001);
ratite birds (Cooper et al. 2001); passerine birds (Erickson et al. 2002);
ichthyophiid caecilians (Gower et al. 2002); and Crypteroniaceae (Conti et al.
2002) and Melastomataceae (Morley and Dick, 2003) plants based on molecular
phylogenetic studies. If these groups of animals and plants reached Asia by
rafting the Indian plate, one would expect to find their fossils in the Cretaceous
- Early Eocene deposits of India. At present there is very limited fossil evidence
for Out-of-India dispersal hypothesis. Myobatrachinae frogs to which
Indobatrachus from the Upper Cretaceous Deccan intertrappean beds of
Bombay is referred today inhabit Australia and Papua-New Guinea region.
A few frog ilia from the intertrappean beds of Naskal assigned to Ranoidea
(Prasad and Rage, 2004) provide the second fossil evidence for the Out-of-
India dispersal hypothesis (Fig.8a-b). Recently, Liu et al. (2007) cited crepe
218 ASHOK SAHNI AND G. V. R. PRASAD

Fig.8a-b. Right ilium (VPL/JU/1013) of Ranoidea frog from the Upper Cretaceous intertrappean
beds of Naskal in lateral (a) and posterior (b) views (after Prasad and Rage, 2004).

myrtle (Lagerstroemia, Lythraceae) from the Deccan intertrappean beds of

India in favour of “Out-of-India” dispersal. In a latest report, Prasad and Bajpai
(2008) documented the presence of agamid lizards from the Early Eocene
Cambay Shale of Vastan Lignite Mine. As mentioned earlier, molecular studies
favoured Gondwanan origin and dispersal to Asia across the rafting Indian
plate for agamid lizards (Macey et al.2000). The presence of oldest agamid
lizards in the Triassic and Jurassic of India and their occurrence in the Early
Eocene of India support molecular data favouring Out-of-India of dispersal for
the agamid lizards.

CONCLUSIONS
The foregoing review clearly demonstrates that the northward flight of
the Indian plate had an important bearing on the evolution of constituent
populations and its biogeographic history. Contrary to expected development
of endemic biota alone during India’s northward drift as a Noah’s Ark, the
fossil record from the Jurassic and Cretaceous deposits indicated the existence
of Gondwanan and Laurasian links in addition to endemic evolution of some
taxa. A longer terrestrial contact between Indo-Madagascar block and South
America (at least as late as 80 Ma) was also inferred based on the fossil record.
The pre-collision (Early Eocene) and collision (Middle Eocene) fauna consists
of endemics as well as Holarctic biotic elements. The Cretaceous and Early
Eocene Holarctic biota may have dispersed to India via island-arc systems as
in the case of Ghazij fauna of Pakistan (Gingerich et al.1997). Currently, the
fossil evidence for Out-of-India dispersal hypothesis is limited. Recent
discoveries of oldest grasses (Prasad et al. 2005), oldest freshwater diatoms
(Ambwani et al. 2003), oldest agamid lizards (Datta and Ray, 2005), supposed
oldest ungulate-like mammal (Prasad et al. 2007b), besides the occurrence of
 GEODYNAMIC EVOLUTION OF THE INDIAN PLATE 219

mammals of archontan affinity (Prasad and Godinot, 1994), oldest lagomorphs

(Rose et al. 2008), and oldest whales (Thewissen et al. 2007) from the Indian
subcontinent added a new dimension to the biodiversity of the Indian plate
during its northward flight. These taxa hint at India as a possible centre of
origin for these groups. However, to test all these propositions, the fossil record
from the Early Cretaceous rocks needs to be improved and search for fossils
from the Palaeocene rocks which have remained elusive until now, needs to be
undertaken in ernst.

ACKNOWLEDGEMENTS
A.S. is thankful to the Indian National Science Academy for Grant No.SP/
SS/2006/2841. G.V.R.P acknowledges financial support from the Department
of Science and Technology, New Delhi (Grant No.SR/S4/ES/24/2002).

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