Vertebrate biodiversity of the Deccan volcanic province of India: A review

Abstract

The Deccan Traps of peninsular India, representing one of the largest flood basalt eruptions on the earth's surface, have
been a subject of intensive research in the last three decades because of the attributed link between the Deccan Traps and
the Cretaceous-Tertiary boundary mass extinctions. In this context, the biota from the sedimentary beds intercalated with
the volcanic flows and underlying the oldest volcanic flow are more important for understanding the faunal diversity and palaeobiogeography
of India during the time span of volcanic eruptions. A detailed review of the vertebrate faunal diversity of the Deccan volcanic
province is presented here.

Full text

doi:10.1130/2014.2505(09)
2014 2014;505; 193-211 , originally published onlineAugust 21,Geological Society of America Special Papers
Guntupalli V.R. Prasad and Ashok Sahni
activity
Vertebrate fauna from the Deccan volcanic province: Response to volcanic
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193
The Geological Society of America
Special Paper 505
2014
Vertebrate fauna from the Deccan volcanic province:
Response to volcanic activity
Guntupalli V.R. Prasad*
Department of Geology, Centre for Advanced Studies, University of Delhi, Delhi 110007, India
Ashok Sahni
98, Mahatma Gandhi Marg, Lucknow 226001, India
ABSTRACT
During the last two decades, extensive paleontological research in the main Dec-
can volcanic province has led to a better understanding of biodiversity close to the
Cretaceous-Paleogene boundary. Several infratrappean localities exposed in Jabal-
pur, Kheda, Balasinor, Rahioli, Dohad, and Bagh in the Narmada Valley (India) pre-
serve one of the most geographically widespread dinosaur nesting sites known in the
world. The well-studied intertrappean beds, such as those of Naskal on the southern
margin, Asifabad and Nagpur on the eastern margin, Kisalpuri and Mohgaon Kalan
on the northeastern margin, and Anjar on the northwestern margin of the main
Deccan volcanic province, have yielded Maastrichtian fi sh (Igdabatis) and dinosaur
remains and palynofossils (Aquilapollenites-Gabonisporites-Ariadnaesporites), either
separately or in association, that suggest a Maastrichtian age for these beds. Only
two intertrappean sections, Papro on the northern margin and Jhilmili on the north-
eastern margin of the main Deccan volcanic province, have produced Paleocene fos-
sils. The fossil record from the infratrappean and intertrappean beds demonstrates
that the dinosaurs survived the early phase of volcanism, though there was an appar-
ent decline in their diversity, and that freshwater vertebrate fauna was least affected
by the initial volcanic activity. The episodic nature of Deccan volcanism may possi-
bly explain the survival of many freshwater and terrestrial communities during the
periods of quiescence. In addition, as in the case of the late Maastrichtian sections
in eastern Montana, North America, detritus-feeding freshwater vertebrate commu-
nities possibly had greater potential for survival than the terrestrial communities
dependent on primary productivity. A close examination of the vertebrate faunal dis-
tribution across the two stratigraphic intervals (infratrappean and intertrappean)
suggests that sampling bias in the infratrappean beds may have also masked the
actual diversity of these beds.
*guntupalli.vrprasad@gmail.com
Prasad, G.V.R., and Sahni, A., 2014, Vertebrate fauna from the Deccan volcanic province: Response to volcanic activity, in Keller, G., and Kerr, A.C., eds., Volca-
nism, Impacts, and Mass Extinctions: Causes and Effects: Geological Society of America Special Paper 505, p. 193–211, doi:10.1130/2014.2505(09). For permis-
sion to copy, contact editing@geosociety.org. © 2014 The Geological Society of America. All rights reserved.
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194 Prasad and Sahni
INTRODUCTION
The Deccan Traps, representing one of the largest continental
fl ood basalts on Earth’s surface, cover about two thirds of penin-
sular India. At many places on the eastern, southern, northern, and
southwestern margins of the main Deccan volcanic province, the
Deccan volcanic fl ows are found in association with sedimentary
beds that have been designated as infratrappean beds (or Lameta
Formation), intertrappean, and supratrappean beds, depending on
their physical position with respect to the volcanic fl ows (Fig. 1).
The infratrappean beds (= Lameta Formation) occur below the
local basal fl ows, whereas the intertrappean beds, deposited dur-
ing the dormant stages of the volcanic activity, occur intercalated
within the lava fl ows. The sediments immediately overlying the
youngest volcanic fl ow are referred as supratrappean beds and
are generally not encountered in outcrops.
Recent radiometric and paleomagnetic studies of basalt fl ows
in the Western Ghats indicate that Deccan volcanism took place in
three short phases interspersed with quiescent periods of consider-
able time (Chenet et al., 2007, 2008, 2009). Phase I, with an esti-
mated volume of ~6%, occurred close to base of C30n at ca. 67.4 Ma
in the late Maastrichtian (Chenet et al., 2007, 2009). Phase II, con-
sisting of several eruptive events of considerable volume (~80%
volume of total Deccan Traps), took place within C29r below the
Cretaceous-Paleogene transition over a short period of time rang-
ing from thousands to tens of thousands of years (Chenet et al.,
2008; Jay and Widdowson, 2008; Jay et al., 2009). Phase III erup-
tions, representing ~14% of Deccan Traps volume, occurred at or
near the base of C29n within the early Danian (Jay and Widdow-
son, 2008; Jay et al., 2009; Keller et al., 2012).
Currently, there are two competing and widely debated
hypotheses to explain the cause of mass extinction at the
Cretaceous-Paleogene (also known as KTB after Cretaceous-
Tertiary) boundary: (1) the asteroid impact hypothesis of Alvarez
et al. (1980) and (2) the volcanic hypothesis of McLean (1985),
Courtillot et al. (1986), and Offi cer et al. (1987). Ever since the
Deccan volcanic activity of India was suggested as a causal link
for this extinction event (McLean, 1985; Offi cer et al., 1987;
Courtillot et al., 1986, 1988), the sedimentary beds occurring in
association with the Deccan Traps (infratrappean, intertrappean,
and supratrappean beds) and yielding plant and animal fossils
have received wide attention from the geoscientifi c community
worldwide. In the last three decades, application of bulk screen-
washing techniques on the infratrappean and intertrappean beds
to recover fossils from different stratigraphic levels of the main
Deccan volcanic province has yielded very promising results. A
variety of vertebrate microfossils, ostracods, molluscs, foramini-
fers, and charophytes previously unknown from the main Dec-
can volcanic province have been documented. In the following
sections, we present a brief history of paleontological research
in the main Deccan volcanic province, the distribution, deposi-
tional environment, and age of infratrappean and intertrappean
beds, and fi nally an analysis of the response of vertebrate fauna
to volcanic activity.
HISTORY OF PALEONTOLOGICAL RESEARCH IN
THE MAIN DECCAN VOLCANIC PROVINCE
There are three phases during which paleontological
research was carried out on the Deccan infratrappean and inter-
trappean beds. During the fi rst phase, which spans India’s pre-
independence era, most of the early studies were carried out by
British geologists, army men, and medical offi cers. W.H. Slee-
man made the fi rst discovery of fossils from the main Deccan
volcanic province in 1828 (Sleeman, 1844). These fossils, rep-
resented by caudal vertebrae, were later referred to a sauropod
dinosaur Titanosaurus indicus by Lydekker (1877), which is now
regarded as an invalid species (Wilson and Upchurch, 2003). Fol-
lowing this, Hislop (1860) published a detailed account on the
molluscan fauna of the Deccan infratrappean and intertrappean
beds. Later on, Woodward (1908) documented the presence of
fi shes (Lepisosteus indicus, Eoserranus hislopi, Pycnodus lam-
etae) from the infratrappean beds of Dongargaon in Chandrapur
District, Maharashtra State (Fig. 2). Additional fi sh remains rep-
resented mainly by scales were described by Hora (1938) from
the intertrappean beds of Deothan and Kheri in Madhya Pradesh
(Fig. 3). Many skull bones, dentitions, and postcranial bones
of dinosaurs were also reported from the infratrappean beds of
Figure 1. Schematic diagram showing
the physical position of infratrappean,
intertrappean, and supratrappean beds
with respect to Deccan volcanic fl ows
(modifi ed after Sahni et al., 1994).
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Vertebrate fauna from the Deccan volcanic province: Response to volcanic activity 195
Figure 2. Map showing the infratrappean vertebrate fossil sites along the Narmada Valley and in Central and South
India. Inset shows outline map of India highlighting the states in which the fossil sites are located.
Figure 3. Map showing vertebrate fossil–yielding intertrappean sites of the Deccan volcanic province. The inset map
of India highlights the states in which these fossil sites are located.
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196 Prasad and Sahni
Pisdura and Dongargaon in Maharashtra, and Jabalpur (Fig. 2)
in Madhya Pradesh (Matley, 1921, 1923, 1929; von Huene and
Matley, 1933). During this phase, frog remains (Indobatrachus
pusillus [Owen, 1847] Noble, 1930) and a turtle plastron (Carter-
emys leithii [Carter, 1852]) were reported from the intertrappean
beds of Bombay (Fig. 3).
In the second phase, representing the period immediately
after the independence of India in 1947 to ca. 1980, the major-
ity of the studies on fossil biota from the main Deccan volcanic
province were confi ned to foraminifers from the intertrappean
beds of Rajahmundry (Bhalla, 1967) or to plant fossils from the
intertrappean beds of Central and Southern India (Prakash, 1960;
Shivarudrappa, 1976–1977; Bhatia and Mannikeri, 1976; Bande
et al., 1986). Some vertebrate fossils, such as frogs, turtle, and
dinosaur remains, were also recorded from the infratrappean
beds of Pisdura, Dongargaon, and Jabalpur (Jain, 1977; Chatter-
jee, 1978) and the intertrappean beds of Bombay (Chiplonkar,
1940; Verma, 1965). Prior to 1980, the Deccan Traps were con-
sidered to span a duration of 30–50 m.y., based on K-Ar dating of
the basaltic fl ows (Alexander, 1981) and molluscs, charophytes,
and other plant fossils from the intertrappean beds (Hislop, 1860;
Sahni, 1934; Hora, 1938; Prakash, 1960; Bhatia and Mannikeri,
1976; Shivarudrappa, 1976–1977).
The third phase of research on the Deccan infratrappean and
intertrappean biota started in the beginning of 1980. Following
the publications by McLean (1985) linking Deccan Traps to the
Cretaceous-Paleogene boundary mass extinction and Courtillot
et al.’s (1986) interpretation of Deccan volcanism as a short-
duration event (<1.0 m.y.) close to the Cretaceous-Paleogene
boundary, renewed interest was generated in the study of Deccan
infratrappean and intertrappean biota. Moreover, the introduction
of bulk screen-washing methods in the early 1980s for the collec-
tion of vertebrate microfossils substantially increased the diver-
sity of various vertebrate groups. Prior to this phase of research
activity, very few vertebrate taxa were known from the Lameta
Formation and intertrappean beds of eruptive phase II. Follow-
ing the application of bulk screen-washing techniques, almost
all vertebrate groups (fi shes, amphibians, turtle, snakes, lizards,
turtles, crocodiles, dinosaurs, and mammals), with the exception
of birds, have been reported from the infratrappean and intertrap-
pean beds (Table 1). In addition to vertebrate microfossils, ~100
species of ostracods have also been documented from the infrat-
rappean and intertrappean beds during phases II and III (see Kho-
sla et al. [2011] for references), and planktic foraminifera were
discovered in the intertrappean beds between phase II and phase
III basalt fl ows at Jhilmili and Rajahmundry (Fig. 3; Keller et al.,
2008, 2009a, 2009b).
DISTRIBUTION, DEPOSITIONAL ENVIRONMENT,
AND AGE OF THE INFRATRAPPEAN AND
INTERTRAPPEAN BEDS
The infratrappean beds and phase II and phase III intertrap-
pean beds are widely known throughout the main Deccan vol-
canic province in outcrops and subsurface sections, whereas the
supratrappean beds are known only from subsurface sections.
Though marine infratrappean and intertrappean strata spanning
the Cretaceous-Paleogene boundary have been delineated in
more than 50 Oil and Natural Gas Corporation (ONGC) deep
wells in Krishna-Godavari Basin (Raju et al., 1991; Jaiprakash et
al., 1993; Keller et al., 2011, 2012), the present paper is restricted
to predominantly continental outcrops from the main Deccan
volcanic province.
Infratrappean Beds or Lameta Formation
Distribution
The infratrappean beds are exposed in discontinuous patches
from Balasinor (Gujarat) in the west to Ambikapur (Madhya
Pradesh) in the east, and from Sagar (Madhya Pradesh) in the
north to Dongargaon, Pisdura (Maharashtra), and Marepalli
(Andhra Pradesh) in the south (Fig. 2), and they reach a maxi-
mum thickness of 75 m, with a thick section known from the type
area at Jhiraghat near Jabalpur in Central India. In the type sec-
tion near Jabalpur, this formation consists of a Green Sandstone,
Lower Limestone, Mottled Nodular Bed, Upper Limestone, and
Upper Sandstone (Matley, 1921). The latter two units were redes-
ignated as Upper (calcifi ed) Sandstone by Tandon et al. (1995).
Not all these units are preserved in the western part and in the
Nand-Dongargaon Basin, where the Upper (calcifi ed) Sandstone
occurs as the most common lithologic unit.
Depositional Environment
The infratrappean beds (Lameta Formation) consist of
sandstones, clays, limestones, and marls that are underlain by
Precambrian basement or rocks of the Gondwana Supergroup.
From time to time, the infratrappean beds have been interpreted
as continental (Matley, 1921; Sahni and Mehrotra, 1974), fl uvial
and shallow-marine (Kumar and Tandon, 1979), shallow-marine
(Chanda, 1967), tidal and estuarine (Singh and Srivastava, 1981;
Singh, 1981; Saha et al., 2010), and pedogenically modifi ed
palustrine mudfl at deposits (Brookfi eld and Sahni, 1987). Sedi-
mentological and isotopic analyses of pedogenic carbonates and
calcretes favored an alluvial-plain setting, with braided streams,
palustrine fl ats, and sheetfl oods under semiarid climatic condi-
tions for the Lameta outcrops of Jabalpur (Ghosh et al., 1995;
Tandon et al., 1995). The Lameta Formation of the Kheda and
Dongargaon areas was interpreted as pedogenically altered over-
bank, channel, paludal, and lacustrine deposits of an alluvial-
plain setting (Mohabey, 1996). Oxygen and carbon isotope stud-
ies of sauropod dinosaur eggshells from the infratrappean beds
of Balasinor led Sarkar et al. (1991) to conclude that the Lameta
dinosaurs consumed water from freshwater pools in a semiarid
climate and ate C3 plants. On the whole, the burden of evidence,
both paleontological and sedimentological, favors a pedogeni-
cally altered, paludal/lacustrine depositional environment in an
alluvial-plain setting under semiarid climate conditions during
deposition of the Lameta Formation.
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Vertebrate fauna from the Deccan volcanic province: Response to volcanic activity 197
TABLE 1. LIST OF VERTEBRATE FOSSILS DESCRIBED FROM THE DECCAN INFRA- AND INTERTRAPPEAN BEDS
Stratigraphic position Taxon Locality State
Intertrappean beds phase II volcanism
Fishes:
Igdabatis indicus Prasad and Cappetta, 1993 Asifabad, Naskal Andhra Pradesh
Gurmatkal Karnataka
Na
g
pur Maharashtra
Kisalpuri Madh
y
a Pradesh
Piplanara
y
anwar Madh
y
a Pradesh
Rhombodus sp. Asifabad Andhra Pradesh
Piplanara
y
anwar Madh
y
a Pradesh
Raja sudhakari Prasad and Cappetta, 1993 Asifabad Andhra Pradesh
Rajiforme indet. (Prasad and Cappetta, 1993) Asifabad Andhra Pradesh
Siluriformes indet. (Cione and Prasad, 2002) Naskal Andhra Pradesh
Clupeidae (Hora, 1938; Prasad and Srinivasan, 1990) Gurmatkal Karnataka
Deothan-Kheri Madh
y
a Pradesh
C
y
prinidae (Hora, 1938) Deothan-Kheri Madh
y
a Pradesh
Pol
y
canthidae (Hora, 1938) Deothan-Kheri Madh
y
a Pradesh
Serranidae (Hora, 1938 Deothan-Kheri Madh
y
a Pradesh
Nandidae (Hora, 1938) Deothan-Kheri Madh
y
a Pradesh
Gurmatkal Karnataka
Pristolepidae (Hora, 1938) Deothan-Kheri Madh
y
a Pradesh
Gurmatkal Karnataka
Horaclupea intertrappea Borkar, 1973 Bamanbor Gujarat
Palaeopristolepis feddeni Borkar, 1973 Bamanbor Gujarat
Palaeopristolepis chiplonkari Borkar, 1984 Bamanbor Gujarat
Indiaichth
y
s bamanborensis Arratia et al., 2004 Bamanbor Gujarat
Percomorpha indet. (Arratia et al., 2004) Bamanbor Gujarat
P
y
cnodontidae indet. Naskal, Asifabad Andhra Pradesh
Rajahmundr
y
Andhra Pradesh
Na
g
pur Maharashtra
Piplanara
y
anwar Madh
y
a Pradesh
Gurmatkal Karnataka
Lepidotes sp. (Mohabey, 1996) Nagpur Maharashtra
Lepisosteus indicus Woodward, 1908 Na
g
pur Maharashtra
Piplanara
y
anwar Madh
y
a Pradesh
Kisalpuri Madh
y
a Pradesh
Gujri Gate Madh
y
a Pradesh
Naskal, Asifabad Andhra Pradesh
?Belonostomus sp. (Ga
y
et et al., 1984) Na
g
pur Maharashtra
Phareodus sp., Osteo
g
lossidae indet. Naskal, Asifabad Andhra Pradesh
Nagpur, Kelapur Maharashtra
Kisalpuri Madh
y
a Pradesh
Piplanara
y
anwar Madh
y
a Pradesh
Deothan-Kheri Madh
y
a Pradesh
Enchodus ferox Leid
y
, 1855 Na
g
pur Maharashtra
Enchodus sp. (Ga
y
et et al., 1984) Na
g
pur Maharashtra
Palaeolabrus cf. dormaalensis (Prasad and Sahni, 1987) Asifabad Andhra Pradesh
Palaeolabrus sp. (Ga
y
et et al., 1984) Na
g
pur Maharashtra
?Sph
y
raena (Ga
y
et et al., 1984) Na
g
pur Maharashtra
Percoidei indet. (Gayet et al., 1984) Nagpur Maharashtra
Ostracion sp. (Ga
y
et et al., 1984) Na
g
pur Maharashtra
Otoliths:
Heterotidinarum heterotoides Nolf et al., 2008 Naskal Andhra Pradesh
Osteo
g
lossidarum deccanensis Rana, 1988 Naskal, Ran
g
apur Andhra Pradesh
Osteo
g
lossidarum intertrappus Rana, 1988 Naskal, Ran
g
apur Andhra Pradesh
Notopteridarum nolfi Rana, 1988 Naskal, Ran
g
apur Andhra Pradesh
Cheemala
g
utta Andhra Pradesh
Clupeidarum valdiyai Rana and Sahni, 1989 Nagpur Maharashtra
Clupeidarum sahnii (Rana, 1996) Ran
g
apur Andhra Pradesh
Gonor
y
nchidarum rectan
g
ulus (Rana, 1988) Naskal, Ran
g
apur Andhra Pradesh
Na
g
pur Maharashtra
Gonor
y
nchidarum sp. (Nolf et al., 2008) Ran
g
apur Andhra Pradesh
Na
g
pur Maharashtra
?Ariidae (Nolf et al., 2008) Naskal, Ran
g
apur Andhra Pradesh
Ariidae indet. (Bajpai and Srinivasan, 1996) Anjar Gujarat
Anthracoperca bhatiai (Rana, 1996) Naskal, Ran
g
apur Andhra Pradesh
Percoideorum citreum Nolf et al., 2008 Naskal Andhra Pradesh
Percoideorum na
g
purensis (Rana and Sahni, 1989) Na
g
pur Maharashtra
Percoideorum ran
g
apurensis Rana, 1988 Naskal, Ran
g
apur Andhra Pradesh
(Continued)
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198 Prasad and Sahni
Percoidei sp. 1 (Nolf et al., 2008) Naskal, Ran
g
apur Andhra Pradesh
Percoidei sp. 2 (Nolf et al., 2008) Ran
g
apur Andhra Pradesh
Percoidei indet. (Bajpai and Srinivasan, 1996) Anjar Gujarat
?Centropomidarum takliensis (Rana and Sahni, 1989) Nagpur Maharashtra
Dapalis erici Nolf et al., 2008 Ran
g
apur Andhra Pradesh
Ambassidarum cappettai (Rana and Sahni, 1989) Na
g
pur Maharashtra
Apo
g
onidarum curvatus (Rana, 1996) Naskal, Ran
g
apur Andhra Pradesh
?Pristolepidinarum jae
g
eri (Rana and Sahni, 1989) Na
g
pur Maharashtra
Blenniidarum sp. (Nolf et al., 2008) Ran
g
apur Andhra Pradesh
Serranidarum sp. 1 (Bajpai and Srinivasan, 1996) Anjar Gujarat
Serranidarum sp. 2 (Bajpai and Srinivasan, 1996) Anjar Gujarat
Amphibians:
Indobatracus pusillus (Owen, 1847) Noble, 1930 Bombay Maharashtra
Gobiatinae (Disco
g
lossidae, ?H
y
lidae) Naskal Andhra Pradesh
Ranoidea (Ranidae or Rhacophoridae) (Prasad and Ra
g
e, 2004). Naskal Andhra Pradesh
Leptodact
y
lidae or Hemisotidae (Prasad and Ra
g
e, 2004) Naskal Andhra Pradesh
Kisalpuri Madh
y
a Pradesh
Pelobatidae indet. (Sahni et al., 1982) Na
g
pur Maharashtra
Lizards:
An
g
uidae indet. (Prasad and Ra
g
e, 1995) Naskal Andhra Pradesh
Litakis sp. (Rana, 2005) Nagpur Maharashtra
Pristi
g
uana sp. (Rana, 2005) Na
g
pur Maharashtra
I
g
uanidae indet. (Rana, 2005) Na
g
pur Maharashtra
A
g
ama sp. (Rana, 2005) Ran
g
apur Andhra Pradesh
?Conto
g
en
y
s sp. (Rana, 2005) Na
g
pur Maharashtra
Eumeces sp. (Rana, 2005) Ran
g
apur Andhra Pradesh
Exostinus estesai Rana, 2005 Na
g
pur Maharashtra
Gekkonid e
gg
shells (Sahni et al., 1984) Na
g
pur Maharashtra
Anjar Gujarat
Turtles:
Carterem
y
s leithii (Carter, 1852) Williams, 1953 Bomba
y
Maharashtra
cf. Carterem
y
s leithii (de Lapperent de Broin et al., 2009) Lakshmipur Gujarat
Carterem
y
s sp.(de Lapperent de Broin et al., 2009) Kisalpuri Madh
y
a Pradesh
Sankuchem
y
s sethnai Gaffne
y
et al., 2003 Bomba
y
Maharashtra
Cf. Bothrem
y
didae (de Lapperent de Broin et al., 2009) Upparhatti Karnataka
Crocodiles:
Crocod
y
lia indet. (Rana, 1990; Prasad and de Lapparent de Broin, 2002) Naskal, Asifabad Andhra Pradesh
Nagpur, Kelapur Maharashtra
Kisalpuri, Madh
y
a Pradesh
Piplanara
y
anwar Madh
y
a Pradesh
Gujri Gate Madh
y
a Pradesh
Anjar Gujarat
Bomba
y
Maharashtra
Crocod
y
lidae indet. (Sin
g
h et al., 1998) Bomba
y
Maharashtra
D
y
rosauridae indet. (Khosla et al., 2009a) Kisalpuri Madh
y
a Pradesh
E
gg
shells (Sin
g
h et al., 1998). Bomba
y
Maharashtra
Snakes:
Indophis sahnii Ra
g
e and Prasad, 1992 Naskal Andhra Pradesh
Kelapur Maharashtra
Anjar Gujarat
Gujri Gate Madh
y
a Pradesh
Piplanara
y
anwar Madh
y
a Pradesh
Madtsoiidae indet. (Ra
g
e et al., 2004) Kelapur Maharashtra
Serpentes incertae sedis (Ra
g
e and Prasad, 1992) Naskal Andhra Pradesh
Coniophis sp. (Rage et al., 2004) Naskal Andhra Pradesh
Dinosaurs:
Indeterminate sauropod bones (Rao and Yadagiri, 1981; Ghevariya, 1988;
Mathur and Sharma, 1990)
Asifabad Andhra Pradesh
Ranipur Madh
y
a Pradesh
Anjar Gujarat
Theropod teeth (Massospond
y
lus rawesi, L
y
dekker, 1890) Na
g
pur Maharashtra
Anjar Gujarat
TABLE 1. LIST OF VERTEBRATE FOSSILS DESCRIBED FROM THE DECCAN INFRA- AND INTERTRAPPEAN BEDS (Continued)
Stratigraphic position Taxon Locality State
Intertrappean beds phase II volcanism
(Continued)
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Vertebrate fauna from the Deccan volcanic province: Response to volcanic activity 199
Gujri Gate Madh
y
a Pradesh
Asifabad Andhra Pradesh
Sauropod e
gg
shells (Sahni et al., 1984) Asifabad Andhra Pradesh
Nagpur, Kelapur Maharashtra
Piplanara
y
anwar Madh
y
a Pradesh
?H
y
pselosaurus sp. (Srinivasan, 1996) Moh
g
aon Kalan Madh
y
a Pradesh
Subtiliolithus kachchhensis (Khosla and Sahni, 1995) Anjar Gujarat
Ornithoid e
gg
shells (Bajpai et al., 1993) Anjar Gujarat
Gujri Gate Madh
y
a Pradesh
Kisalpuri Madh
y
a Pradesh
Mammals:
Deccanolestes hislopi Prasad and Sahni, 1988 Naskal, Ran
g
apur Andhra Pradesh
Deccanolestes robustus Prasad et al., 1994 Naskal, Ran
g
apur Andhra Pradesh
Deccanolestes narmadensis Prasad et al., 2010 Kisalpuri Madh
y
a Pradesh
Sahnitherium ran
g
apurensis Rana and Wilson, 2003 Ran
g
apur Andhra Pradesh
Kharmerun
g
ulatum vanvaleni Prasad et al., 2007a Kisalpuri Madh
y
a Pradesh
Bharattherium bonapartei Prasad et al., 2007b (=Dakshina jederi Wilson et
al., 2007)
Kisalpuri Madhya Pradesh
Naskal Andhra Pradesh
Infratrappean beds below phase I and/or phase II volcanism
Fishes:
I
g
dabatis indicus Prasad and Cappetta, 1993 Pisdura Maharashtra
Jabalpur Madh
y
a Pradesh
Marepalli Andhra Pradesh
Nand-Don
g
ar
g
aon Maharashtra
Rhombodus sp. Marepalli Andhra Pradesh
Lepisosteus indicus Woodward, 1908 Dongargaon Maharashtra
Marepalli Andhra Pradesh
Jabalpur Madh
y
a Pradesh
P
y
cnodus lametae Woodward, 1908 Don
g
ar
g
aon Maharashtra
P
y
cnodontidae indet. Jabalpur Madh
y
a Pradesh
Eoserranus hislopi Woodward, 1908 Don
g
ar
g
aon Maharashtra
Lepidotes deccanensis S
y
kes, 1851 Nand-Don
g
ar
g
aon Maharashtra
Clupea sp. (Mohabe
y
, 1996) Nand-Don
g
ar
g
aon Maharashtra
Nandidae indet. (Mohabe
y
, 1996) Marepalli Andhra Pradesh
Osteo
g
lossidae indet. (Mohabe
y
, 1996) Marepalli Andhra Pradesh
Don
g
ar
g
aon Maharashtra
Jabalpur Madhya Pradesh
Enchodus sp. (Jain and Sahni, 1983; Mohabe
y
, 1996) Nand-Don
g
ar
g
aon Maharashtra
Pisdura Maharashtra
Marepalli Andhra Pradesh
Arius sp. (Jain and Sahni, 1983) Pisdura Maharashtra
Crocodiles:
Crocod
y
lia indet. Marepalli Andhra Pradesh
D
y
rosauridae indet (Rana, 1987) Auspalli, Andhra Pradesh
Marepalli Andhra Pradesh
Turtles:
Shweboem
y
s pisdurensis (Jain, 1986) Pisdura Maharashtra
E
gg
shells (Bajpai et al., 1997) Rajahmundr
y
Andhra Pradesh
Snakes:
Sanajeh indicus (Madtsoiidae) Wilson et al., 2010 Kheda Gujarat
Madtsoia pisdurensis (Madtsoiidae) Mohabe
y
et al., 2011 Pisdura Maharashtra
Dinosaurs:
Jainosaurus (=Antarctosaurus) septentrionalis (von Huene and Matley, 1933)
Hunt et al., 1994
Pisdura Maharashtra
Jabalpur Madhya Pradesh
Jainosaurus cf. septentrionalis (Wilson et al., 2011) Jabalpur Madh
y
a Pradesh
Isisaurus colberti (Jain and Bandyopadhyay, 1997) Wilson and Upchurch,
2003
Dongargaon Maharashtra
Titanosauriformes indet. (Wilson and Mohabe
y
, 2006) Nand Maharashtra
Indosuchus raptorius von Huene and Matle
y
, 1933 Jabalpur Madh
y
a Pradesh
Indosaurus matleyi von Huene and Matley, 1933 Jabalpur Madhya Pradesh
Laevisuchus indicus von Huene and Matle
y
, 1933 Jabalpur Madh
y
a Pradesh
Lametasaurus indicus von Huene and Matle
y
, 1933 Jabalpur Madh
y
a Pradesh
TABLE 1. LIST OF VERTEBRATE FOSSILS DESCRIBED FROM THE DECCAN INFRA- AND INTERTRAPPEAN BEDS (Continued)
Stratigraphic position Taxon Locality State
Intertrappean beds phase II volcanism
(Continued)
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200 Prasad and Sahni
Age
For a long time, the Lameta Formation was regarded as
Turonian in age, based on the similarity of dinosaur fauna
between the Lameta Formation and the Maevarano Formation of
Madagascar, which was initially assigned a Turonian age (von
Huene and Matley, 1933; Chiplonkar, 1986). While reviewing the
Lameta dinosaur fauna, Buffetaut (1987) rejected the Turonian
age for the Lameta Formation and favored a Maastrichtian age.
The latter age was confi rmed by the subsequent discoveries of a
Maastrichtian fi sh (Igdabatis, Rhombodus; Jain and Sahni, 1983;
Courtillot et al., 1986) and a palynological assemblage consisting
of Ariadnaesporites and Aquilapollenites (Dogra et al., 1988).
The Maastrichtian age for the Lameta Formation was further
supported by a 40Ar/39Ar age of 66.4 ± 1.9 Ma (Courtillot et al.,
1988) for the basal fl ow overlying the Maastrichtian sediments
at Dongargaon (Fig. 2). However, paleomagnetic studies of the
volcanic fl ows overlying the Lameta Formation at Jabalpur had
placed these fl ows in magnetochron C30n (Courtillot et al., 1986)
and thus in phase I of Deccan volcanism. Hence, the fossils and
radiometric data from the Lameta Formation underlying the basal
volcanic fl ows indicate a late Maastrichtian lower age limit for
the Deccan volcanic eruptions.
Intertrappean Beds
Distribution
The intertrappean sedimentary beds are found mainly along
the southern, eastern, northeastern, and northwestern margins of
the main Deccan volcanic province. The well-known intertrappean
sections are from Gurmatkal and Upparhatti (Karnataka State),
Naskal, Rangapur, and Asifabad (Andhra Pradesh State), Nag-
pur and Bombay (Maharashtra State), Mohgaon Kalan, Jhilmili,
Kisalpuri, Mandla, Padwar, Ranipur, Piplanarayanwar, and Gujri
Gate (Madhya Pradesh State), Anjar and Bamanbor (Gujarat
State), Mamoni (Rajasthan State), and Papro (Uttar Pradesh state)
(Fig. 3). At present, there are no paleomagnetic or geochronologi-
cal controls on the position of the intertrappean beds between the
lava fl ows. However, based on geochemical characteristics, Jay
and Widdowson (2008) correlated the basaltic fl ows on the south-
eastern, eastern, northeastern, and northern margins of the main
Deccan volcanic province with the Poladpur and Ambenali For-
mations of the Wai Group in the Western Ghats. In view of this
and the presence of Maastrichtian fossils and dinosaur remains in
the intertrappean beds of Gurmatkal, Naskal, Rangapur, Asifabad,
Nagpur, Mohgaon Kalan, Kisalpuri, Mandla, Padwar, Ranipur,
Piplanarayanwar, Gujri Gate, Anjar, and Mamoni (Fig. 3), we
prefer to place these beds in phase II volcanism.
The intertrappean beds of Jhilmili (Fig. 3), with early Paleo-
cene fossils and lower and upper traps placed within chrons C29r
and C29r/C29n, respectively, were considered to lie between
phases II and III (Keller et al., 2009b). The intertrappean beds
of Papro (Fig. 3), with Paleocene palynofossils, may also be
placed between volcanic phases II and III. The intertrappean beds
of Bombay were placed within phase III (Cripps et al., 2005).
Currently, there are limited fossil data to pinpoint the position
of Upparhatti and Bamanbor (Fig. 3) intertrappean beds within
the lava fl ow stratigraphy of the main Deccan volcanic province.
Compared with the Lameta Formation, the intertrappean beds of
phases II and III are relatively thin, generally reaching a few cen-
timeters to 30 cm in thickness, and rarely measuring up to 5 m,
except at Jhilmili, where these intertrappean beds measure 13.5 m
(Keller et al., 2009a, 2009b).
Depositional Environment
Since the pioneering work of Hislop (1860), many intertrap-
pean outcrops have been investigated from the paleontological
point of view. Of these, the intertrappean beds of phase II, such
as Rangapur, Naskal, Asifabad, Nagpur, Kisalpuri, Piplanaray-
anwar, and Anjar (Fig. 3), have been intensively studied for ver-
tebrate microfossils. Generally, they are composed of siltstones,
Rajasaurus narmadensis Wilson et al., 2003 Rahioli Gujarat
Rahiolisaurus
g
ujaratensis Novas et al., 2010 Rahioli Gujarat
Oospecies:
Me
g
aloolithus c
y
lindricus Khosla and Sahni, 1995 Jabalpur Madh
y
a Pradesh
M. mohabe
y
i Khosla and Sahni, 1995 Dholi
y
a Madh
y
a Pradesh
M. padi
y
alensis Khosla and Sahni, 1995 Padi
y
al Madh
y
a Pradesh
M. jabalpurensis Khosla and Sahni, 1995 Jabalpur Madh
y
a Pradesh
M. dholi
y
aensis Khosla and Sahni, 1995 Dholi
y
a Madh
y
a Pradesh
M. dhoridun
g
riensis Mohabe
y
, 1998 Dhoridun
g
ri Gujarat
M. khempurensis Mohabe
y
, 1998 Khempur Gujarat
M. me
g
adermus Mohabe
y
, 1998 Dohad Gujarat
M. ba
g
hensis Khosla and Sahni, 1995 Ba
g
h Madh
y
a Pradesh
Sauropod eggshells (Jain and Sahni, 1985) Pisdura Maharashtra
Ellipsoolithus khedaensis (Family Elongatoolithidae) (Mohabey, 1998; Loyal
et al., 1998).
Kheda Gujarat
Mammals:
Avashishta bacharamensis Anantharaman et al., 2006 Bacharam Andhra Pradesh
TABLE 1. LIST OF VERTEBRATE FOSSILS DESCRIBED FROM THE DECCAN INFRA- AND INTERTRAPPEAN BEDS (Continued)
Stratigraphic position Taxon Locality State
Infratrappean beds below phase I
and/or phase II volcanism
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Vertebrate fauna from the Deccan volcanic province: Response to volcanic activity 201
mudstones, sandstones, clays, marls, and shales, frequently asso-
ciated with a prominent fossiliferous chert and occasionally with
a limestone bed. The Deccan intertrappean beds are traditionally
considered freshwater lacustrine deposits that fi lled low-lying
areas on the surface of the lava fl ows during the dormant stages
of volcanism.
The phase II intertrappean beds of Naskal and Anjar (Fig. 3)
were studied in some detail in the context of their depositional
environment. Based on sedimentological, paleontological, and
taphonomic studies, the Naskal intertrappean beds were inter-
preted as alkaline lake deposits in a fl oodplain setting intermit-
tently subjected to subaerial exposure, leading to the develop-
ment of two phases of paleosols (Khajuria and Prasad, 1998).
Sedimentological studies of the iridium-bearing Anjar intertrap-
pean section indicated a freshwater lacustrine environment in a
semiarid climate setting (Khadkikar et al., 1999). However, the
mixed presence of marine and nonmarine biota in the intertrap-
pean beds of phase II at Asifabad and Nagpur (Fig. 3) indicates
some marine infl uence in these areas (Sahni, 1983; Prasad and
Sahni, 1987). This has been proven by the recent discovery of
planktonic foraminifers from the intertrappean beds occurring
between phase II and phase III basalt fl ows at Jhilmili in Central
India (Keller et al., 2009b).
Age
Early paleontological studies overestimated the age of
Deccan intertrappean beds as Early Tertiary (with the majority
favoring an Eocene age) based on fi sh, molluscs, and plant fos-
sils, including charophytes and plant megafossils (Hislop, 1860;
Sahni, 1934; Hora, 1938; Bhatia and Mannikeri, 1976; Shivaru-
drappa, 1976–1977; Bande et al., 1986), and the absence of dino-
saurs as compared to their frequent occurrence in the Lameta
Formation (Matley, 1921, 1929; von Huene and Matley, 1933,
Chatterjee, 1978). Khajuria et al. (1994) discussed in detail the
inadequacy of these fossils for assigning an Eocene age for the
intertrappean beds. Further, dinosaur remains (a few bones and
teeth, and several eggshell fragments) have been found in many
intertrappean beds (Rao and Yadagiri, 1981; Vianey-Liaud et al.,
1987; Prasad, 1989; Sahni and Bajpai, 1988; Ghevariya, 1988;
Mathur and Sharma, 1990).
Subsequent fi ndings of Maastrichtian fi shes (Prasad and
Cappetta, 1993) and the Ariadnaesporites-Aquilapollenites-
Gabonisporites palynological assemblage (Mathur and Sharma,
1990; Sahni et al., 1996; Kumaran et al., 1997; Dogra et al., 2004;
Singh et al., 2006) from the majority of the intertrappean beds
favor a Maastrichtian age. A couple of intertrappean outcrops
from the main Deccan volcanic province have been assigned a
younger early Paleocene age. The northernmost intertrappean
section near Papro in Lalitpur District (Uttar Pradesh State) has
been assigned an early Paleocene age based on a palynological
assemblage consisting of Dandotiaspora dilata, Dandotiaspora
pseudoauriculata, Dandotiaspora plicata, Spinizonocolpites
echinatus, Matanomadhiasulcites sp., and Lakiapollis ovatus
(Singh and Kar, 2002).
The Anjar intertrappean section in Kachchh, western India,
was initially designated as Cretaceous-Paleogene age based on
anomalous concentrations of iridium in the sediments sandwiched
between volcanic fl ows III and IV (Bhandari et al., 1996). How-
ever, later paleontological studies demonstrated that the inter-
trappean beds overlying the iridium-bearing levels yield Maas-
trichtian fossils, including dinosaur remains (Bajpai and Prasad,
2000), and the high iridium concentration was attributed to leach-
ing from the Deccan volcanic fl ows (Khadkikar et al., 1999). As
a result, the iridium-enriched levels of the Anjar intertrappean
beds are no longer regarded as representing the Cretaceous-
Paleogene boundary. Therefore, no fossiliferous intertrappean
outcrop spanning the Cretaceous-Paleogene transition has been
recorded from the main Deccan volcanic province to date.
More recently, Keller et al. (2009a, 2009b) reported the pres-
ence of early Danian planktic foraminifers in the intertrappean
beds between volcanic phases II and III at Jhilmili in Chhindwara
District (Madhya Pradesh State). However, the Maastrichtian to
Danian transition has yet to be discovered in this area.
On the whole, the fossil record from the infratrappean and
intertrappean outcrops is in agreement with the fossil record
from the subsurface intertrappean sections of ONGC wells in the
Krishna-Godavari Basin, where the shallow-marine foraminiferal
assemblages from the infratrappean beds and phase II intertrappean
beds indicate a late Maastrichtian age, and phase III intertrappean
and supratrappean beds indicate an early Paleocene (Danian) age
for the Deccan volcanism (Raju et al., 1991; Jaiprakash et al., 1993;
Keller et al., 2011, 2012). The paleontological data from infratrap-
pean and intertrappean beds from the main Deccan volcanic prov-
ince outcrops and from shallow-marine subsurface sections are in
agreement with radiometric dates from a thick Deccan volcanic
pile from the Western Ghats, which indicate that Deccan volcanism
began ca. 67.4 Ma and terminated around 62 Ma (Venkatesan et al.,
1993; Chenet et al., 2007, 2008).
RESPONSE OF VERTEBRATE FAUNA TO
DECCAN VOLCANISM
One frequently asked question is whether Deccan volcanism
was capable of affecting the ecosystems at the global level, and,
if so, what were its effects at the site of eruption? To seek answers
to these questions, one should examine the fossil evidence from
the sediments deposited prior to the initiation of volcanism (infra-
trappean) and those deposited during (intertrappean) or immedi-
ately after Deccan volcanic activity.
Fishes, Crocodiles, Turtles, and Snakes
A comparison of the infratrappean and intertrappean verte-
brate fauna demonstrates that at least 10 infratrappean fi sh taxa
(fi ve each at species and family level) survived into the inter-
trappean beds of phase II (Table 1; Fig. 4). Although some inde-
terminate anurans and lizards were reported by Jain and Sahni
(1983) and Mohabey (1996), no recognizable amphibian and
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202 Prasad and Sahni
lizard remains have been recorded to date from the infratrap-
pean beds. On the other hand, the intertrappean amphibians and
lizards are represented by at least four and six families, respec-
tively. Two crocodilian groups (Crocodylia indet. and Dyrosau-
ridae) of the infratrappean beds survived into the intertrappean
beds (Fig. 4). The lone turtle species (Shweboemys pisdurensis
Jain, 1986) known from the infratrappean beds belongs to the
family Podocnemididae. In comparison, the intertrappean turtles
are identifi ed as Carteremys leithii (Carter, 1852), Sankuchemys
sethnai Gaffney et al., 2003 of the family Bothremydidae, and an
indeterminate form distinct from all these taxa (de Lapparent de
Broin et al., 2009). The infratrappean snake fauna is represented
Figure 4. Range chart of various vertebrate groups across the Deccan volcano-sedimentary sequences.
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Vertebrate fauna from the Deccan volcanic province: Response to volcanic activity 203
by two taxa: Sanajeh indicus Wilson et al., 2010 and Madtsoia
pisdurensis Mohabey et al., 2011 of the family Madtsoiidae. Not
only did the madtsoiid snakes survive into the intertrappean beds
at the family level, but so did three new taxa: Indophis sahnii
(?Nigerophiidae) Rage and Prasad, 1992, Serpentes incertae
sedis (Rage and Prasad, 1992), and Coniophis (Aniliidae) (Rage
et al., 2004) appear at this stratigraphic level.
Dinosaur Bones, Nesting Sites, Eggshells, and Teeth
Lameta Formation
One of the most important fossil fi ndings from the main Dec-
can volcanic province in the last three decades is the discovery
of geographically extensive nesting sites and eggshells in the
Lameta Formation (below phase II volcanism) over a stretch of
1000 km from Jabalpur in the east to Balasinor in the west and
in the Nand-Dongargaon Basin in Central India (Fig. 2). These
nesting sites are located near Jabalpur in the upper Narmada Val-
ley, Bagh, Jobat, Dohad, Kheda, Balasinor, and Rahioli in the
lower Narmada Valley, and Pavna and Dongargaon in the Nand-
Dongargaon Basin (Fig. 2). In all these sites, nests and eggshells
belonging to nine oospecies of Megaoolithus (Table 1) were doc-
umented from a fi ne- to medium-grained calcretized sandstone
(Srivastava et al., 1986; Mohabey, 1998; Vianey-Liaud et al.,
2003). Only one oospecies of Ellipsoolithus (Table 1) has so far
been described from the Lameta Formation near Kheda (Moha-
bey, 1998). The occurrence of nests in a similar lithofacies has
been attributed to strong nesting site selectivity, and extensiveness
of nesting sites with morphologically similar eggs was shown as
evidence for colonial nesting (Sahni et al., 1994; Mohabey, 2001).
Besides nesting sites and eggshells, the Lameta Formation
at Jabalpur, Balasinor, Pisdura, Dongargaon, and Nand has been
the source for all known Cretaceous dinosaur taxa from India
named on the basis of skull and postcranial bones (Sleeman,
1844; Lydekker, 1877; Matley, 1921, 1923, 1929; von Huene
and Matley, 1933; Chatterjee, 1978; Berman and Jain, 1982;
Srivastava et al., 1986; Mathur and Srivastava, 1987; Jain and
Bandyopadhyay, 1997; Wilson et al., 2003; Novas et al., 2010;
Wilson and Mohabey, 2006; Wilson et al., 2011). To date, three
titanosaurid sauropods and six abelisaurid theropods have been
described from the Lameta Formation.1
Intertrappean Beds
Another signifi cant fi nding from the main Deccan volcanic
province is the occurrence of dinosaur remains in the intertrap-
pean beds. All the early reports of skeletal remains of dinosaurs,
coprolites, and dinosaur nesting sites were from the Lameta For-
mation (Matley, 1921; Matley, 1929; von Huene and Matley,
1933; Chatterjee, 1978; Berman and Jain, 1982; Srivastava et al.,
1986; Mathur and Srivastava, 1987). Prior to 1980s, dinosaurs
were considered to have been restricted to the Lameta Formation,
and the intertrappean beds were generally regarded as lacking
dinosaur fossils. This was one of the reasons for assigning an
Early Tertiary (Eocene) age for the intertrappean beds. However,
since 1980, several dinosaur fossils, including a few bones and
teeth, and many eggshell fragments, have been documented from
phase II intertrappean beds. Moreover, they have been reported
from widely separated intertrappean sections such as those of
Asifabad (Rao and Yadagiri, 1981; Sahni et al., 1984; Prasad,
1989), Nagpur (Vianey-Liaud et al., 1987), Anjar (Ghevariya,
1988; Sahni and Bajpai, 1988; Bajpai et al., 1993; Bajpai and
Prasad, 2000), Ranipur (Mathur and Sharma, 1990), Mohgaon
Kalan (Srinivasan, 1996), and Kisalpuri (Khosla et al., 2004)
(Fig. 3). More recently, dinosaur eggshell fragments have also
been found in the intertrappean beds of Kelapur (Maharashtra)
and Piplanarayanwar and Gujri Gate (Madhya Pradesh) (Fig. 3)
(work in progress at Delhi University).
These intertrappean eggshells are generally thin as com-
pared to those of the Lameta Formation. Although they have
not been studied as extensively and in as much detail as those
of the Lameta Formation, limited ultrastructural studies indi-
cate that eggshells from Asifabad, Nagpur (Sahni et al., 1984;
Vianey-Liaud et al., 1987), Mohgaon Kalan (Srinivasan, 1996),
Kelapur, and Piplanarayanwar belong to sauropod dinosaurs,
and those from Anjar (Bajpai et al., 1993), Kisalpuri, and Gujri
Gate are assignable to the ornithoid type. Until now, no paratax-
onomic classifi cation has been attempted for the intertrappean
dinosaur eggshells.
Depositional Environment of Dinosaur Fossils: Lameta
versus Intertrappean
The Lameta Formation is generally thick, reaching a maxi-
mum thickness of 75 m, and the dinosaur nests and eggs are pref-
erentially preserved in a sandy calcretized unit of the carbonate
facies at Jabalpur, Rahioli, and Dongargaon (Mohabey, 2001).
At Jabalpur, the dinosaur skeletal remains were recovered from
the base and within the lower limestone. At Rahioli, dinosaur
bones occur within a conglomerate horizon underlying the dino-
saur egg-bearing sandy calcrete (Mohabey, 2001). In the Nand-
Dongargaon Basin, dinosaur bones are reworked and transported
elements within the overbank red and green clays, whereas the
eggs are exclusively found in sandy calcrete and channel sand-
stone (Mohabey, 2001). In the area near the Bagh Caves in the
lower Narmada Valley, dinosaur-egg yielding levels of the Lam-
eta Formation are confi ned to the sandy and nodular calcretes of
massive cherty limestone (Mohabey, 2001).
In contrast, the intertrappean beds are relatively thin (gen-
erally a few centimeters to 30 cm thickness, rarely reaching 5
m) with widely varying lithologies consisting of soft siltstone,
1Titanosaurid sauropods: Jainosaurus (= Antarctosaurus) septentrionalis (von
Huene and Matley, 1933) Hunt et al., 1994; Jainosaurus cf. septentriona-
lis (Wilson et al., 2011); Isisaurus colberti (Jain and Bandyopadhyay, 1997)
Wilson and Upchurch, 2003; Titanosauriformes indet. (Wilson and Mohabey,
2006). Abelisaurid theropods: Indosuchus raptorius von Huene and Matley,
1933; Indosaurus matleyi von Huene and Matley, 1933; Laevisuchus indicus
von Huene and Matley, 1933; Lametasaurus indicus von Huene and Matley,
1933; Rajasaurus narmadensis Wilson et al., 2003; and Rahiolisaurus gujara-
tensis (Novas et al., 2010) (see Table 1).
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204 Prasad and Sahni
shale, mudstone, marl, and clay. Sporadic dinosaur bones have
been reported from the phase II intertrappean beds at Asifabad,
near Hyderabad (Rao and Yadagiri, 1981), and Ranipur (Mathur
and Sharma, 1990). However, the comparatively thick (~3.5 m)
intertrappean beds of Anjar have yielded several large, yet to be
described dinosaur bones (Ghevariya, 1988).
A major difference between the dinosaur fauna of the Lam-
eta Formation and phase II intertrappean beds is the abundance of
bones and nesting sites and fully preserved eggs in the former as
compared to relatively limited presence of bones and no complete
eggs or nesting sites in the latter. In view of the fact that the Lam-
eta and phase II intertrappean dinosaur fossils were preserved in
two distinctly different lithofacies, it is diffi cult to say whether
there was a true decline of dinosaurs from the Lameta Formation
to phase II intertrappean beds or whether this is an artifact of
preservation. Depositional environments seem to have played an
important role in the preservation of dinosaur fossils. The hard,
sandy carbonate facies of the Lameta Formation deposited in a
coastal-plain environment and proximal to the dinosaur habitat
might have ensured better preservation of bones and complete
eggs. In comparison, the intertrappean depositional environments
represented by thin lacustrine facies distal to the dinosaur habitat
may have received transported eggshell fragments and occasional
large bones.
Mammals
At least two major groups of mammals have been docu-
mented from three widely separated phase II intertrappean sites.
The intertrappean beds of Naskal near Hyderabad (Fig. 3) yielded
the most diversifi ed microvertebrate assemblage, including the
fi rst Cretaceous mammals of India. Deccanolestes hislopi was
the fi rst eutherian mammal recorded from the intertrappean beds
of Naskal (Prasad and Sahni, 1988). Following this report, a few
more taxa, such as Deccanolestes robustus (Prasad et al., 1994)
and Sahnitherium rangapurensis (Rana and Wilson, 2003), were
also documented from this area. Later on, one more species of
Deccanolestes (Deccanolestes narmadensis Prasad et al., 2010)
and a possible ungulate (Kharmerungulatum vanvaleni Prasad
et al., 2007a) were described from the intertrappean beds of
Kisalpuri in Central India (Fig. 3). Phylogenetic analysis of Dec-
canolestes has shown that it belongs to an insectivorous group of
mammals (Adapisoriculidae), and this group possibly originated
in India and dispersed to Africa and Europe some time close to
the Cretaceous-Paleogene boundary (Goswami et al., 2011; de
Bast et al., 2012).
These discoveries of eutherian mammals from the main
Deccan volcanic province assume great signifi cance in light of
recent molecular phylogenies suggesting the former Gondwana-
land as a center of origin for certain placental mammal orders
(Springer, 1997; Waddell et al., 1999). As compared to a mod-
erately diverse mammalian assemblage known from the inter-
trappean beds, only one possible haramiyidan mammal tooth
(Avashishta bacharamensis Anantharaman et al., 2006) has been
described from the infratrappean beds. The latter was recovered
from the infratrappean beds of Bacharam, Rangareddi District,
Andhra Pradesh (Anantharaman et al., 2006). The poor mam-
malian record from the infratrappean beds can be attributed to
sampling bias. So far, samples from 10 phase II intertrappean
sections have been bulk screen-washed, out of which only three
turned out to be mammal-bearing. Out of four infratrappean sec-
tions (i.e., Pisdura, Jabalpur, Bacharam, and Marepalli; Fig. 2)
subjected to bulk screen-washing, only one has produced a mam-
malian fossil. Thus, future works focused on washing of large
numbers of samples may improve the diversity of infratrappean
mammalian fauna.
SURVIVAL PATTERNS: LAMETA
VERSUS INTERTRAPPEAN
The Lameta (infratrappean) fauna is represented by 12 fi sh,
two crocodile, one turtle, two snake, and nine dinosaur taxa (three
sauropod and six theropod) and 10 dinosaur oospecies (Table 1).
In comparison, the phase II intertrappean vertebrate fauna is rep-
resented by 47 fi sh species (including 21 otolith species), fi ve
amphibian, eight lizard, four snake, three crocodile, four turtle,
and six mammalian taxa. Besides these, sauropod and ornithoid
eggshells, theropod teeth, and a few large sauropod bones are
also known from the intertrappean beds. The diversity of inter-
trappean vertebrate fauna more than doubled, at least in the case
of fi sh, snakes, and turtles at species and higher taxonomic levels.
This diversity increase may be due to sampling bias or the
availability of more nutrients in the ecosystem following the ini-
tial phase of volcanism. Although the infratrappean beds have
been extensively prospected for dinosaurs, very few sections
have been explored for vertebrate microfossils by means of bulk
screen-washing methods. In contrast, large numbers of intertrap-
pean sections have been investigated based on this method. In
addition, following erosion and transportation of materials from
the newly deposited mafi c volcanic rocks, many new nutrients
might have been added to the lacustrine bodies, which may have
played an important role in increasing their biodiversity. To
discriminate between these two plausible explanations for the
increased biodiversity in the intertrappean beds, more infratrap-
pean sections need to be sampled on a larger scale.
Overall, the infratrappean vertebrate fauna survived Deccan
volcanism phase I and, with the exception of dinosaurs, prolifer-
ated in the intertrappean ecosystems. This pattern of vertebrate
survival from the infratrappean to intertrappean beds has a close
parallel with the continental late Maastrichtian record of the
western interior of North America (eastern Montana). There, the
uppermost Cretaceous (late Maastrichtian) Hell Creek Forma-
tion yielded 107 vertebrates species, of which 52 species (49%)
survived into the lower part of the overlying Lower Paleocene
Tullock Formation (Archibald, 2011). Compared to 28% survival
in land-dwelling animals, freshwater animals such as bony fi shes,
amphibians, turtles, crocodiles, and champsosaurs had a high
survival rate of 76%. Sheehan and Fastovsky (1992) inferred a
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Vertebrate fauna from the Deccan volcanic province: Response to volcanic activity 205
higher rate of survival for freshwater faunas (90%) as compared
to land-dwellers (12%). This differential pattern of extinction of
freshwater and terrestrial vertebrates has been explained in terms
of dependence of land-based communities on primary produc-
tivity in contrast to buffering of freshwater communities from
extinction by detritus feeding when there was a temporary drop
in primary productivity (Sheehan and Fastovsky, 1992).
A similar explanation can be offered for the preferential sur-
vival of the predominantly freshwater aquatic community com-
pared with the terrestrial community from the time of deposition
of infratrappeans to intertrappean beds. There is a notable increase
in the diversity of the intertrappean freshwater vertebrates such as
fi shes, turtles, and snakes as compared to that of the infratrappean
vertebrates (Fig. 5). The freshwater vertebrates constitute 52%,
while the terrestrial vertebrates form 49% of the infratrappean
vertebrate fauna (Fig. 5). In contrast, the intertrappean vertebrate
fauna is dominated by freshwater vertebrates (70%), compared to
terrestrial vertebrates (30%). Additionally, it is quite possible that
depositional environments also played a signifi cant role in the
preservation of fossils. The humid, tropical, lacustrine environ-
ments of the intertrappean beds had possibly greater potential for
the preservation of vertebrate fossils as compared to overbank,
paludal environments of the semiarid climate during deposition
of the infratrappean beds. Prasad and Khajuria (1995) suggested
that the survival of Deccan phase I volcanism by freshwater com-
munities might be related to the episodic nature of the volcanic
eruptions, and the fauna could have recovered from the environ-
mental stress during the repose periods.
Invertebrates also seem to have responded to the initial
Deccan volcanic activity in this manner. Khosla et al. (2011)
described 41 species of freshwater ostracods from the Lameta
Formation of Jabalpur, Pisdura, and Dongargaon. According to
these authors, of the 41 species, fi ve are indeterminate, six are
new, and 30 are common to the intertrappean beds. Besides their
common occurrence in Maastrichtian infratrappean and inter-
trappean beds, many of these Maastrichtian species are found
in association with Danian planktic foraminifers and brackish-
water ostracod species in Jhilmili intertrappean beds (Keller et
al., 2009b). This demonstrates that the ostracods of the Maas-
trichtian intertrappean beds survived into the early Paleocene,
apparently without major extinctions due to volcanism (Sharma
and Khosla, 2009; Khosla et al., 2011).
According to Cripps et al. (2005), the third and last phase of
Deccan volcanism did not affect life signifi cantly. Cripps et al.
(2005) considered the intertrappean beds of Bombay as part of the
Mumbai Island Formation (Salsette Subgroup) overlying the Wai
Subgroup of main Deccan volcanic province and hence represen-
tative of the fi nal phase III of Deccan volcanism that followed
the Cretaceous-Paleogene boundary mass extinction. Singh and
Sahni (1996) assigned a Maastrichtian age for the intertrappean
beds of Bombay on the basis of an ostracod assemblage similar
to that of other intertrappean outcrops of the main Deccan vol-
canic province. However, the recent report of similar ostracod
assemblage in association with early Paleocene (Danian) plank-
tic foraminifers from the intertrappean beds of Jhilmili argues
against the use of ostracods as age markers for the intertrappean
beds. Palynofacies analysis of Bombay intertrappean beds led
Cripps et al. (2005) to conclude that these beds were deposited in
a lagoonal environment proximal to land, sheltered from a strong
marine infl uence and more organic rich in comparison to those
of the main Deccan volcanic province. Because of the absence
of evidence for wildfi res that would cause mass mortality and
increase the availability of plant material to the lagoonal ecosys-
tem throughout the numerous pyroclastic eruptions in this region,
Cripps et al. (2005) argued that the late-stage explosive Deccan
volcanism might not have caused a major ecosystem collapse.
However, based on palynofl oral analysis of the Lameta For-
mation and succeeding intertrappean beds in Nand- Dongargaon
Basin in Maharashtra, Samant and Mohabey (2005) concluded
that the initial Deccan volcanic activity affected the plant com-
munity. The fl oral transition from the Lameta Formation to
the overlying intertrappean stratigraphic levels is marked by
a decline in diatom diversity and abundance, an increase in
angiosperm and pteridophyte diversity at the expense of gym-
nosperms, and the fi rst appearance of dinofl agellates (Samant
and Mohabey, this volume). They attributed these fl oral changes
to initial physiographic and chemical changes caused by Dec-
can volcanism. Though Samant and Mohabey (2005) observed
total absence of diatoms and dinofl agellates in spore-pollen–
rich intertrappean beds at higher stratigraphic levels, the spore-
pollen assemblages from the intertrappean beds of different
stratigraphic levels are generally more or less similar. Even
the pollen-spore assemblage represented by Ariadnaesporites,
Aquilapollenites, Gabonisporites, Azolla, etc. is broadly simi-
lar to Lameta and intertrappean beds. A marked difference in
the palynofl oral assemblage has only been noticed between the
Maastrichtian outcrops and the Paleocene intertrappean beds of
Papro, Lalitpur (Singh and Kar, 2002).
SUMMARY
Intensifi ed vertebrate faunal studies in recent years have
revealed the presence of widespread nesting sites over an area
of 1000 km in an east-west direction along the Narmada Valley
and in Central India. Following the application of bulk screen-
washing methods, there has been a remarkable improvement in
the record of diversity of vertebrate fauna of the Deccan infrat-
rappean and intertrappean beds. As a consequence, all vertebrate
groups, with the exception of birds, have been recovered from
the intertrappean beds. In comparison to the common occurrence
of dinosaur skeletal remains and nesting sites and eggs in the
Lameta Formation, the phase II intertrappean beds yielded only
isolated teeth and bones in a few sites and eggshell fragments
at many localities. This apparent decline of the dinosaur fauna
subsequent to the eruption of the fi rst phase of Deccan volca-
nism might be attributed to taphonomic bias, as the infratrappean
and intertrappean dinosaur-bearing sedimentary facies are dis-
tinct from each other. A comparison of vertebrate fauna from the
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206 Prasad and Sahni
Figure 5. Relative composition of
freshwater aquatic and terrestrial ver-
tebrates of the infratrappean and inter-
trappean beds.
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Vertebrate fauna from the Deccan volcanic province: Response to volcanic activity 207
infratrappean and intertrappean beds revealed that many fresh-
water taxa survived the fi rst phase of Deccan volcanism. The rea-
sons one could offer for their survival are: (1) ability of the fauna
to bounce back during the quiescent periods of volcanism; (2) the
different feeding strategies adapted by freshwater and terrestrial
communities; or (3) sampling bias—the observed increase in the
vertebrate diversity from the infratrappean to intertrappean beds
could be a consequence of limited sampling from the infratrap-
pean beds in comparison to large-scale screen-washing of inter-
trappean sediments. If Deccan volcanism had any impact on life,
it was possibly during the second phase, when 80% of the total
volume erupted. Although a linkage has been suggested between
Deccan volcanism and foraminiferal extinction in Cretaceous-
Paleogene boundary subsurface sections of ONGC wells in the
Krishna-Godavari Basin (Keller et al., 2011, 2012) and in the
Um Sohryngkew section of Meghalaya, ~800 km northeast of the
main Deccan volcanic province (Gertsch et al., 2011), so far no
well-defi ned Cretaceous-Paleogene boundary section has been
discovered in continental outcrops of the main Deccan volcanic
province. Therefore, future studies should focus on identifying
clearly marked Cretaceous-Paleogene boundary sections in the
main Deccan volcanic province outcrops in order to understand
the effects of volcanism on physical environments and biota in a
terrestrial environment.
ACKNOWLEDGMENTS
Prasad acknowledges the research grant from J.C. Bose National
Fellowship of Department of Science and Technology, New
Delhi. We are thankful to Gerta Keller for inviting us to contrib-
ute to this volume. This manuscript has benefi ted from critical
reviews by Gerta Keller, W.A. Clemens, and Jeffrey A. Wilson.
Help from L. Ranjit Singh, Department of Geology, University
of Delhi, in making the fi gures is duly acknowledged.
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