Vertebrate

Fauna of the
Cauvery Basin,
Southern India
PRIYA SINGH, and Dr. Moumita Das*
Department of Geology,
Banaras Hindu University ,Varanasi-221005
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ABSTRACT
 Vertebrate fossils have been known from South India’s Cauvery Basin since the 1840s, but
records of marine vertebrates from the late Albian to Turonian Karai Formation have been
limited to a single set of ichthyosaur remains. Recent surface collecting and sieving of lower
Cenomanian glauconitic mudstones has yielded the first ichthyosaur material reported in
India over the last 140 years, as well as a diverse and previously unrecorded shark
assemblage. The ichthyosaur material, including several teeth and vertebrae, is assigned to
the sole described Cretaceous genus Platypterygius and to the species P. indicus (Lydekker,
1879). Eight species of shark (one squaliformes, two hexanchiformes, and five lamniformes)
are recorded. A new hexanchiform genus Gladioserratus is erected, and two new species
(Gladioserratus magnus, gen. et sp. nov., and Dwardius sudindicus, sp. nov.) are named
 The vertebrate fauna from the late Albian to Maastrichtian succession of the Cauvery Basin, south
India has been known since 1845, but it received only scant attention as compared to the vertebrates
already known from the Deccan volcanic province of peninsular India. Recent fossil discoveries
appear to support the emergence of significantly diverse marine and non-marine vertebrates
comprising fishes, frogs, reptiles (ichthyosaurs, plesiosaurs, turtles, crocodiles and dinosaurs) and a
mammal from the Cauvery Basin have revived the interest in the fauna of the basin as it has
significant implications for understanding the palaeobiogeography of India.
 The latest Albian to Turonian marine vertebrates such as sharks, ichthyosaurs and plesiosaurs show
a wide geographic distribution and marine territory, while sharks are typically cooler water fauna of
high palaeolatitudes. The latest Maastrichtian non-marine vertebrates especially turtles, crocodiles,
dinosaurs and a mammal are considered to show mixed Gondwanan and Laurasian affinities thus
providing new lines of evidence in favour of a latest Cretaceous biotic links between India and the
neighbouring continents. An overview of the vertebrate faunal diversity of the Cretaceous sequences
of the Cauvery Basin and its palaeobiogeographic considerations are presented.
INTRODUCTION
The Cretaceous vertebrate fauna of India has significant implications in terms
of evolution and palaeobiogeography because of its position relative to the rest
of Gondwana at a time of dispersal of the Gondwanan continental masses
(Prasad, 2006). The Indian subcontinent thus had a key biogeographic position
between the Eastern Gondwana and Eurasia during the Cretaceous to Eocene.
It was once a part of Gondwana continent and most of the palaeogeographic
reconstructions depict that the Indo-Madagascar block was linked to Western
Gondwana (South America and Africa) during most of the Jurassic (Lawver et
al., 1998). During the Late Jurassic (ca. 160 Ma ago), the Indo-Madagascar
block broke away from Africa and in the Early Cretaceous (ca. 120 to 130
Ma), it dismembered from Australia.
India, separated from Madagascar by ca. 88 Ma ago as an isolated subcontinent (Storey et al., 1995)
and eventually collided with Eurasia in the Palaeocene/ Early Eocene (ca. 50 to 55 Ma) after
following its northwest journey in the middle of the Tethys for a period of more than 30 to 35 Ma
(Barron and Harrison, 1980; Aitchison et al., 2007; Chatterjee et al., 2013). Cretaceous vertebrate
fossils from India and especially those from the Cauvery Basin are thus important to address various
biogeographic issues associated with the movement of the tectonic plates. The record of the
Cretaceous vertebrates of India mainly comes from the Upper Cretaceous sediments. Although India
has well-preserved sedimentary deposits of the Lower Cretaceous age, these units are typically poorly
fossiliferous and have yielded few fossils of palaeobiogeographic significance (Prasad et al., 2006;).
The most thoroughly described of the Late Cretaceous vertebrate faunas of India are those recovered
from the sedimentary successions associated with the Deccan volcanics (Prasad, 2007;) and to some
extent from the Bagh Formation in the Narmada Basin, central western India. These vertebrate faunas
comprise frogs, lizards, snakes, crocodiles, turtles, dinosaurs and mammals and have received
extensive attention as far as the evolution and biogeography of the Late Cretaceous vertebrates of
India are concerned. Despite the fact that Cretaceous deposits of the Cauvery Basin, south India have
been known to yield vertebrate remains since 1845, its vertebrate fauna only received significant
attention recently.
STUDY AREA
DISCUSSION
As the northward drift of India continued, sediments of the Trichinopoly and Ariyalur
groups were deposited in the Ariyalur subbasin of the Cauvery basin. Yadagiri and Ayyasami
(1979) reported a partial skull and postcranial skeleton, attributed to a stegosaur
Dravidosaurus blanfordi, from the Coniacian Anaipadi Formation of the Trichinopoly Group
from a site near Siranattam village. Latter examination of these bones led others to doubt its
identification (Chatterjee and Rudra, 2013) or regard it as nomen dubium or Stegosauria
indet. Vertebrate fossils have been recovered at three levels within the Maastrichtian
interval. A sauropod dinosaur egg belonging to Megaloolithus cylindricus (Ariyalur Group),
originally known from the Upper Cretaceous Lameta Formation of Cental India, was
described from the shallow marine Upper Cretaceous (Lower Maastrichtian)
Kallankuruchchi.
More recently, Dhiman et al. (2017) reported sauropod dinosaur eggshell fragments representing
Fusioolithus baghensis from the shallow marine lower part of the Upper Cretaceous (Upper
Maastrichtian) Kallamedu Formation. Eggshells with microstructure similar to that of M.
cylindricus have been described from the Upper Cretaceous rocks of France and Upper Cretaceous
Allen Formation of Argentina.

Megaloolithus pseudomammilare from the Upper Maastrichtian deposits of Aix-en-Provence,

France and Tremp Basin, Spain and Patagoolithus salitralensis from the Upper Cretaceous of
Salitral Moreno, Argentina were also considered as junior synonyms of F. baghensis. Hence this
sauropod oospecies demonstrate close palaeobiogeographic links with both Southern Hemisphere
continents and southern Europe.
The upper continental part of the Kallamedu Formation is long known for its fragmentary dinosaur
bones.
Large limb and girdle bones comparable to those of titanosaurid sauropods of the Lameta
Formation of Central India were identified in the fossil bones from the Kallamedu Formation
reported bones of theropod, sauropod, and stegosaur dinosaurs from the Kallamedu Formation
and described some of them as representing a new theropod dinosaur Bruhathkayosaurus
matleyi. According to Galton and Ayyasami (2017), the bones of B. matleyi do not exist any
more as they disintegrated in the plaster jackets before reaching Geological Survey of India
headquaters and B. matleyi is now regarded either as nomen dubium or Sauropoda indet.

Following a detailed study of supposed ornithischian dinosaur bone identified as a stegosaur

dermal plate from the Upper Cretaceous Kallamedu Formation. Galton and Ayyasami (2017)
concluded that this bone probably belongs to a sauropod. The Kallamedu Formation yielded
remains of fishes: Lepisosteidae indet., Egertonia sp. amphibian: Anura indet. (Prasad et al.,
2007); turtle: Kurmademys kallamedensis (Gaffney et al., 2001); crocodiles: cf. Simosuchus
sp., Crocodylia indet. (Prasad et al., 2007); dinosaurs: Abelisauridae indet. (Prasad et al.,
2006), Troodontidae indet. and mammals: Sudamericidae indet.

An isolated theropod tooth described from the Kallamedu site as Megalosaurus sp. by
Lydekker (1879) may actually belong to the abelisaurid dinosaurs.
The Kallamedu fauna thus has clades that have pan-Gondwanan affinities
(gondwanatherian mammals and abelisaurid dinosaurs) (Goswami et al., 2012;
Prasad et al., 2013) or Madagascan affinities (Egertonia sp., Kurmademys,
simosuchid crocodyliform( Prasad et al., 2007) on one hand and a clade of Laurasian
affinity such as troodontid theropod dinosaur on the other hand. Halliday et al.
(2017) comparing fauna of all the Late Cretaceous localities of India and
Madagascar demonstrated that the faunal composition of the Kallamedu Formation
is distinct fromthat of the Deccan infra- and intertrappean beds, rather it shows
greater faunal similarity with the Upper Cretaceous Maevarano Formation of
Madagascar, despite the fact that the two landmasses were separated several million
years ago.
Cretaceous vertebrate diversity
Vertebrate remains have long been known from the Cretaceous sequences of the
Cauvery Basin (e.g., Lydekker, 1879), but only little consideration has been given
to understand their diversity and palaeobiogeographic implications to date. Recent
reports of diverse fossil records from the Cauvery Basin are remarkably improved
the knowledge about vertebrate diversities and also added new important forms
( Prasad et al., 2007).

In addition, the new data also allows direct comparisons with contemporaneous
vertebrates known from other continents such as South America, Africa, Madagascar
and Europe and thus leading to infer crucial palaeobiogeographic patterns, in the
context of the latest available palaeocoastline reconstructions of the Gondwanan
continents. The vertebrate fauna from the Cauvery Basin is now known to include
fishes, frogs, reptiles (ichthyosaurs, plesiosaurs, turtles and dinosaurs) and a
mammal.
Fishes
Fishes were the earliest discovered vertebrates from the Cauvery Basin. During the
investigation of Cretaceous fossiliferous sequences in the Pondicherry and adjacent
areas, south India between 1840 and 1842, C.J. Kaye and C.E. Cunliffe collected
fish remains from limestone beds and handed over their collection to Sir Philip
Egerton for systematic studies identified twelve shark species, and one form of
pycnodont and enchodont actinopterygian. Stoliczka (1873) documented the
presence of durophagous shark, Ptychodus latissimus Agassiz, 1843 and
pycnodont,?Pycnodus sp., from the Uttattur Group of Ariyalur sub-basin, while
Blanford (1862) also mentioned the presence of Ptychodus and Pycnodus in the
Uttattur Group. Later, Lydekker (1887) mentioned the occurrence of the same
assemblage in Cretaceous deposits of Ariyalur sub-basin and all these authors
maintained that most of the fishes are closely related to European forms. The
current whereabouts of the fish material described by Egerton (1845) is unknown
and no worker was able to re-examine the material; the species erected by Egerton
(1845) are still lacking a valid taxonomic assignment and hence not mentioned.
However, in the most cases, precise locations and ages of fish-yielding horizons were not provided.
Cretaceous rocks exposed at Pondicherry and Vridhachalam areas where Kaye and Cunliffe seemingly
collected sharks teeth are most likely chronologically as well as lithologically to be coeval to the
Ariyalur Group of the Ariyalur sub-basin. The Cretaceous marine clastic rocks of Pondicherry and
Vridhachalam regions are considered to be Santonian to Maastrichtian in age.
It is, therefore, reasonable to assume that shark remains described from the neighbourhood of
Pondicherry area were collected from the Santonian to Maastrichtian deposits. After a break of nearly
a hundred years, Paul (1973) described a shark tooth, Oxyrhina sp., from the Uttattur Group. More
recently, Underwood et al. (2011) recorded some more marine shark species such as Protosqualus sp.,
Gladioserratus magnus Underwood et al., 2011, Notidanodon sp., Cretalamna appendiculata Agassiz,
1843, Dwardius sudindicus Underwood et al., 2011, ?Eostriatolamia sp., Squalicorax aff. baharijensis
Stromer, 1927 and Cretodus longiplicatus Werner, 1989 from the late Albian to Turonian Karai
Formation. Additionally, Verma described one more species of a durophagous shark, Ptychodus
decurrens Agassiz, 1843, from the Karai Formation (Fig. 3m–n). Prasad et al. (2006) reported isolated
ganoid scales of lepisosteiformes from the freshwater deposits of the Kallamedu Formation. Otolith-
based fish species such as Lycoclupea menakiae Gowda, 1967, belonging to actinopterygian and an
unnamed otolith as well as two ossiculiths described as specimens A and B were also reported from the
Karai Formation
Frogs
The record of amphibians from the Cauvery Basin is almost non-existent as
compared to frogs known from the Upper Cretaceous Deccan volcanic
province. Recently, a fragmentary ilium belonging to an indeterminate
anuran has been documented from the Maastrichtian freshwater sediments
of the Kallamedu Formation (Prasad et al., 2006). This is the only known
report of any frog from the Cauvery Basin.

Reptiles
Marine reptiles are poorly known from the Cauvery Basin. Lydekker (1879) erected a
new species of ichthyosaur, Platypterygius indicus Lydekker, 1879, based on fifteen
complete and fragmentary vertebrae from the Karai Formation.
Recently, Underwood et al. (2011) recovered a few isolated, complete teeth and
partially fragmentary vertebrae, tentatively described as P. indicus. The authors,
however, pointed out that a detailed examination is necessary to validate the taxonomic
status of this species. McGowan and Motani (2003) also proposed that material of P.
indicus is not sufficient to erect a new species and considered the status of P. indicus as
indeterminate. Yadagiri and Ayyasami (1979) collected some skeletal material from
Coniacian deposits of the Anaipadi Formation, Trichinopoly Group on which they
erected a new stegosaur dinosaur, Dravidosaurus blanfordi Yadagiri and Ayyasami,
1979. Subsequent studies of the material doubted on its dinosaur affinity and suggested
that these remains may belong to a plesiosaur .

Sahni (1957) described a fossilized reptilian egg, referred to a chelonian that was
recorded from the Cenomanian sediments of the Uttattur Group.

The presence of fossil turtles in the Uttattur Group was first noted by Muzzy (1857)
and Blanford (1862), but till date no turtle fossil has been fully described from this
group. However, a cast of the interior shell of a turtle was reported from the early
Turonian sediments of the Karai Formation.
In discussing the cast of the turtle, the authors mentioned that the material is not sufficient to be
identified at species level, but observed that the cast of the animal had an elongated shell, elevated in
the vertebral region with sloping sides and had prominent sutural unions of the pelvis. Based on these
observations, the authors indentified it as a testudine pleurodiran turtle. In contrast, quite recently two
fossil turtles from the Kallamedu Formation were reported. Gaffney et al. (2001) described a single
well-preserved, nearly complete skull and some postcranial remains of a turtle and assigned it to a
new genus and species Kurmademys kallamedensis Gaffney et al., 2001. It is a side-necked
pelomedusid bothremydid and the authors also mentioned that the material may include five other
more poorly preserved and unprepared skulls.
Subsequently, Gaffney et al. (2009) proposed that three bothremydids such as Kurmademys from the
Kallamedu, Sankuchemys from the Upper Cretaceous sediments associated with the Deccan
volcanism of Mumbai and Kinkonychelys from the Upper Cretaceous Maevarano Formation of
Madagascar are nested in the tribe Kurmademydini and are sister to each other. In addition, several
isolated, fragmentary carapaces and one vertebral piece recovered from the Kallamedu Formation
were referred to an indeterminate testudine turtle (Prasad et al., 2007).
Despite more than 170 years of collecting vertebrates from the Cauvery Basin, crocodylian remains
were reported only recently from the Kallamedu Formation (Prasad et al., 2006).
A single significant isolated multicuspid tooth referred to a Simosuchus-like notosuchian and three
isolated teeth along with one osteoderm identified as indeterminate Crocodylia have been described
(Prasad et al., 2007. The report of a Simosuchus-like notosuchian from India is interesting as it
represents the first occurrence of a Simosuchus-like crocodile outside the Late Cretaceous of
Madagascar. Simosuchus-like notosuchians are only currently known from the Late Cretaceous of
Madagascar and India (Prasad et al., 2006).
Dinosaurs are a significant biotic component of the Cauvery Basin and have been known since 1862,
when Blanford discovered a single tooth and ill-preserved bones from the Kallamedu bone-bed of the
Kallamedu Formation. Hundreds of dinosaur remains have been collected from the Kallamedu bone-
bed since then, though very few of which have been identified and described.
Lydekker (1879) identified the single isolated tooth of Blanford's collection as a theropod dinosaur
and assigned it to the Megalosaurus sp. From the same bone-bed, Matley (1929) reported large limb-
bones and girdles of sauropods, probably belonging to the Titanosaurus that was earlier recorded from
the Upper Cretaceous Lameta Formation of Jabalpur, Madhya Pradesh, central India and a few small
bones assigned to stegosaurs. In addition, Matley (1929) also observed and mentioned that bones
occurring in the Kallamedu bone-bed were extremely friable and it was very hard to extract any
complete, identifiable bones even after the greatest care was taken. Naryana Rao and Seshachar
(1927) collected additional bones and concluded that these remains might belong to dinosaurs.
Later, Naryana Rao and Rama Rao (1930) jointly studied a well-preserved vertebra from Naryana
Rao and Seshachar's collection and inferred that it represents the anterior part of a reptilian
vertebra, probably belonging to a camarasaur sauropod. While reexamining the collection of
Naryana Rao and Seshachar, Yadagiri et al. (1983) concluded that the collection was probably
mammalian remains from Pleistocene deposits; the occurrence of Pleistocene mammalian remains
around the Kallamedu bone-bed was already earlier mentioned by Matley (1929). An isolated
well-preserved cervical vertebra recovered from the Kallamedu Formation was referred to the
Megalosauridae (Rama Rao, 1932). During the 1980s, geologists from the Geological Survey of
India excavated the Kallamedu area and collected dinosaur material. Subsequently, Yadagiri and
Ayyasami (1989) described a new genus and species of a carnosaur theropod, Bruhatkayosaurus
matleyi Yadagiri and Ayyasami, 1989, based on material consisting of an ilia, an ischium, a femur,
a tibia, a radius and a vertebra. The tibia is 2 m long and femur 0.75 m, which are about 29% and
33% larger than the tibia and femur, respectively, of Argentinosaurus, a titanosaur sauropod from
the Late Cretaceous of Argentina, indicating that B. matleyi was larger than Argentinosaurus and
is one of the most gigantic dinosaurs ever discovered. However, some authors recently casted
doubt on the taxonomic assignment of B. matleyi to a theropod dinosaur considered B. matleyi as
a sauropod dinosaur
Solitary mammal
A single tooth of a sudamericid gondwanatherian mammal has been recovered
recently from the Kallamedu Formation (Goswami et al., 2012; Prasad, 2013).
Gondwanatheria is an extinct group of non tribosphenic mammals that are reported
only from the Gondwanan continents, particularly South America, Antarctica,
Africa, Madagascar and India,

Earlier, sudamericid mammals were known to occur in the Upper Cretaceous

Deccan intertrappean beds of Madhya Pradesh, Telangana and Karnataka (Prasad et
al., 2007)and their presence in the Cauvery Basin represents the southernmost
extension of sudamericids in India .
However, other groups of mammals such as adapisoriculids, archaic ungulates and
haramiyids known from the Deccan volcanic province are yet to be documented
from the Cauvery Basin (Prasad et al., 2006, 2007 )
CONCLUSION
An overview of the vertebrates from Cretaceous (Albian to Maastrichtian) successions of
the Cauvery Basin, Tamil Nadu, southern India shows that the Uttattur and Trichinopoly
groups, ranging from the Albian to Coniacian, yielded mainly marine vertebrates and the
Maastrichtian Kallamedu Formation of the Ariyalur Group is known to yield non-marine
vertebrates. The marine vertebrate assemblages consist of sharks (Squalicorax, Dwardius,
Cretalamna, ?Eostriatolamia, Protosqualus, Gladioserratus, ?Notidanodon, Cretodus and
Ptychodus), pycnodonts (Pycnodus), enchodonts, otolith-based actinopterygian
(Lycoclupea menakiae), ichthyosaurs (Platypterygius indicus), plesiosaurs (Dravidosaurus
blanfordi), testudine turtles and a reptilian egg shell. Many taxa show cosmopolitan
distributions having broader biogeographical affinities to Laurasian continents, and to some
extent with Gondwana. The reports of pycnodont fishes (Pycnodus and “Coelodus”) from
the shallow marine Uttattur Group and the Bagh Formation extends the Indian record of
pycnodonts from the late Maastrichtian to the latest Albian and Coniacian and it might
have been possible that Maastrichtian pycnodonts from the Deccan volcanic province were
the immigrants of pycnodonts of the Cauvery Basin and the Bagh Formation.
The non-marine vertebrate assemblage comprises frogs, turtles (Kurmademys
kallamedensis), crocodiles (Simosuchus-like notosuchian, Crocodylia indet.),
dinosaurs (Megalosaurus sp., Titanosaurus sp., B. matleyi, Megaloolithus,
Abelisauridae indet. and Troodontidae indet.) and a mammal (Sudamericidae indet.).
The Kurmademys turtle and Simosuchus-like crocodile are suggestive of close
biogeographic connections between India and Madagascar during the latest
Cretaceous. The abelisaurid dinosaurs and sudamericid gondwanatherian mammal
support faunal similarities during the Late Cretaceous between South America and
Indo-Madagascar. The presence of a troodontid dinosaur in the Late Cretaceous of
India and an abelisaurid dinosaur in France may indicate sporadic dispersals
between India and Europe via Africa. Though abelisaurid and troodontid dinosaurs
were present in the Late Cretaceous of France and India, respectively, the
abelisaurids were clearly the more dominant components of Gondwanan continents
particularly, South America, Africa, Madagascar and India whereas the troodontids
were the more dominant faunal elements of Laurasian continents such as Asia,
Europe and North America
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