cretaceous vertebrate fauna of the cauvery basin, southern india: palaeodiversity and...

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Palaeogeography, Palaeoclimatology, Palaeoecology 431 (2015) 53–67

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Cretaceous vertebrate fauna of the Cauvery Basin, southern India:Palaeodiversity and palaeobiogeographic implications

Omkar Verma ⁎Geology Discipline Group, School of Sciences, Indira Gandhi National Open University, New Delhi 110068, India

⁎ Tel.: +91 29531675; fax: +91 29532167.E-mail address: [email protected].

http://dx.doi.org/10.1016/j.palaeo.2015.04.0210031-0182/© 2015 Elsevier B.V. All rights reserved.

a b s t r a c t
a r t i c l e i n f o
Article history:Received 6 November 2014Received in revised form 6 April 2015Accepted 21 April 2015Available online 30 April 2015

Keywords:AlbianMaastrichtianVertebratesPalaeobiogeographyCauvery BasinIndia

The vertebrate fauna from the late Albian toMaastrichtian succession of the Cauvery Basin, south India has beenknown since 1845, but it received only scant attention as compared to the vertebrates already known from theDeccan volcanic province of peninsular India. Recent fossil discoveries appear to support the emergence of signif-icantly 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 faunaof the basin as it has significant implications for understanding the palaeobiogeography of India. The latest Albianto Turonianmarine vertebrates such as sharks, ichthyosaurs and plesiosaurs show awide geographic distributionand marine territory, while sharks are typically cooler water fauna of high palaeolatitudes. The latestMaastrichtian non-marine vertebrates especially turtles, crocodiles, dinosaurs and a mammal are considered toshow mixed Gondwanan and Laurasian affinities thus providing new lines of evidence in favour of a latest Creta-ceous biotic links between India and the neighbouring continents. An overview of the vertebrate faunal diversityof the Cretaceous sequences of the Cauvery Basin and its palaeobiogeographic considerations are presented.

© 2015 Elsevier B.V. All rights reserved.

1. Introduction

The Cretaceous vertebrate fauna of India has significant implicationsin terms of evolution and palaeobiogeography because of its positionrelative to the rest of Gondwana at a time of dispersal of theGondwanancontinental masses (Prasad, 2005). The Indian subcontinent thus had akey biogeographic position between the Eastern Gondwana and Eurasiaduring the Cretaceous to Eocene (Sahni, 2010). It was once a part ofGondwana continent andmost of the palaeogeographic reconstructionsdepict that the Indo-Madagascar blockwas linked toWesternGondwana(South America and Africa) during most of the Jurassic (Lawver et al.,1998). During the Late Jurassic (ca. 160 Ma ago), the Indo-Madagascarblock broke away from Africa (Coffin and Rabinowitz, 1987) and in theEarly Cretaceous (ca. 120 to 130 Ma), it dismembered from Australia–Antarctica (Powell et al., 1988; Chand et al., 2001). India, separatedfrom Madagascar by ca. 88 Ma ago as an isolated subcontinent (Storeyet al., 1995) and eventually collided with Eurasia in the Palaeocene/Early Eocene (ca. 50 to 55 Ma) after following its northwest journey inthe middle of the Tethys for a period of more than 30 to 35 Ma (Barronand 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 biogeographicissues associated with the movement of the tectonic plates.

The record of the Cretaceous vertebrates of Indiamainly comes fromthe Upper Cretaceous sediments (Prasad, 2012). Although India haswell-preserved sedimentary deposits of the Lower Cretaceous age,these units are typically poorly fossiliferous and have yielded few fossilsof palaeobiogeographic significance (Baksi, 1973; Prasad et al., 2004;Tripathi et al., 2013). The most thoroughly described of the LateCretaceous vertebrate faunas of India are those recovered from the sed-imentary successions associated with the Deccan volcanics (Khosla andSahni, 2003; Verma, 2008; Prasad, 2012; Khosla and Verma, 2014;Prasad and Sahni, 2014) and to some extent from the Bagh Formationin the Narmada Basin, central western India (Verma, 1965; Chiplonkarand Ghare, 1977; Khosla et al., 2003). These vertebrate faunas comprisefrogs, lizards, snakes, crocodiles, turtles, dinosaurs and mammals andhave received extensive attention as far as the evolution and biogeogra-phy of the Late Cretaceous vertebrates of India are concerned (Khoslaand Verma, 2014 and references therein). Despite the fact thatCretaceous deposits of the Cauvery Basin, south India have beenknown to yield vertebrate remains since 1845, its vertebrate faunaonly received significant attention recently (Underwood et al., 2011;Verma et al., 2012a; Goswami et al., 2013; Prasad et al., 2013). This com-munication is intended to review the Cretaceous vertebrate diversity ofthe Cauvery Basin and to provide an account of its biostratigraphic,

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Fig. 1. a. Map of India showing the Cauvery Basin. b. Geological map of the Cauvery Basin highlighting three sub-basins viz., Pondicherry, Vridhachalam and Ariyalur (mapmodified afterSastri and Raiverman, 1968). c. Geological map of the Cauvery Basin exposed at the Ariyalur area showing vertebrate fossil-bearing sites (map modified after Sundaram et al., 2001).

54 O. Verma / Palaeogeography, Palaeoclimatology, Palaeoecology 431 (2015) 53–67


55O. Verma / Palaeogeography, Palaeoclimatology, Palaeoecology 431 (2015) 53–67


palaeoenvironmental and palaeobiogeographic implications in the lightof recent findings.

2. Geology, biostratigraphy and palaeoenvironments

The Cauvery Basin is a NE–SW trending rift basin that was formedduring the Late Jurassic to Early Cretaceous splitting of the Archaeanbasement between India and Australia–Antarctica (Veevers et al.,1991). It is a large basin with an approximate aerial extension of25,000 km2, consisting of well-preserved shallow marine sedimentarysequences of the Albian to Maastrichtian age, deposited along the east-ern coast of the Tamil Nadu state, southern India (Sundaramet al., 2001;Fig. 1a). The sedimentary sequences in the Cauvery Basin occur in fiveisolated sub-basins (Fig. 1b) viz., Pondicherry, Vridhachalam, Ariyalur,Tanjore and Sivaganga (Banerji, 1972) with the Ariyalur sub-basin con-taining the most extensive and well-exposed Cretaceous to Palaeocenesuccession (Fig. 1c). Blanford (1862) first studied and mapped theAriyalur area and classified the sequence into three groups: Uttattur,Trichinopoly and Ariyalur that have been differentiated based onlithology and fossils. Following this author, several other workers(e.g., Banerji, 1972; Sastry et al., 1972; Sundaram and Rao, 1986;Ramasamy and Banerji, 1991; Tewari et al., 1996; Sundaram et al.,2001) presented detailed accounts of the lithostratigraphic classifica-tion for the Ariyalur sub-basin. However, the revised version of thelithostratigraphy of this sub-basin presented by Sundaram et al.(2001) retained the original threefold lithostratigraphic subdivision ofthe sub-basin that was initially proposed by Blanford (1862). The latestlithostratigraphic classification and terminology proposed by Sundaramet al. (2001) is shown in Table 1.

The Uttattur Group contains a basal transgressive unit of thesequence, which rests unconformably on the Archaean basement andattains a maximum thickness of about 820 m. The group is subdividedinto four formations viz., Terani, Arogyapurum, Dalmiapuram andKarai in chronological order (Sundaram et al., 2001). The basal TeraniFormation marks the beginning of first Cretaceous sedimentation inthe basin (Ramkumar et al., 2004). It consists of boulder conglomerates,coarse and poorly bedded sandstones, claystones, shales and siltstones.It yielded a rich assemblage of megaplant fossils such as Ptilophyllum,but is poor in megainvertebrates. The dominance of conglomerate andsandstone is considered to be indicative of a high-energy fluvial to del-taic depositional environment (Tewari et al., 1996; Sundaram et al.,2001). Mamgain et al. (1973) documented two Barremian ammonitesand one inoceramid species along with plant fossils from clayey beds.An Albian age has been assigned to the formation by Sundaram et al.(2001). The Arogyapurum Formation comprisesmainly the conglomer-ates andmedium to very coarse grained sandstones. The sedimentologyand geometry of the sandstone bodies indicate that they representfluvial systems. This formation also lacks age diagnostic fossils and isconsidered to be of Albian age based on its stratigraphic position

Table 1Lithostratigraphic classification of the Cretaceous sequence of the Cauvery Basin(after Sundaram et al., 2001).

Group Formation Depositionalenvironments

Age

Ariyalur Kallamedu Fluvial Late MaastrichtianKallankurichchi Shallow marine Early MaastrichtianSillakkudi Littoral to shallow

marineCampanian

Trichinopoly Anaipadi Neritic Turonian to ConiacianKulakkalnattam Littoral to shallow

marineTuronian

Uttattar Karai Shallow marine Late Albian to earlyTuronian

Dalmiapuram Shallow marine AlbianArogyapurum Fluvial AlbianTerani Fluvial to deltaic Albian

(Sundaram et al., 2001). Lithologically, the Dalmiapuram Formationcontains grey shales, coral algal limestones, bedded limestones, pebbleconglomerates, siltstones and marls. It is highly fossiliferous and con-tains abundant remains of corals andmolluscs. The coral algal limestonein places associated with grey shales and the occurrence of pebbleconglomerates in the formation represents a marine flooding and theabundance of benthic foraminifer such as Lenticulina spp., suggestssome 30 m palaeowater depth (Nagendra et al., 2014). It has yieldedammonites indicative of the Mortoniceras inflatum and Mortonicerasrostratum zones and the oyster Rastellum (Arcostrea) carinata, suggest-ing a late Albian age (Sastry et al., 1968; Ayyasami, 2006). However,the presence of foraminifers such as Hedbergella planispira PartialRange Zone, Rotalipora appenninica Interval Zone to R. cushmani TotalRange Zone, Ticinella roberti Total Range Zone and the belemnitesTetrabelus seclusus, Parahebolites blanfordi and Neohibolites sp. favour amiddle Albian to late Cenomanian age range (Ayyasami and Rao,1987; Tewari et al., 1996; Sundaram et al., 2001). The uppermostKarai Formation contains clays, silty clays, sandy clays, siltstones, sand-stones and occasionally phosphatic nodules and gypsum. The formationis further subdivided into two members, the lower Odiyam andthe upper Kunnam members. The Odiyam Member is argillaceous,composed of clay with abundant phosphatic nodules and gypisiferousmudstone and occasionally intercalated with sandstone and siltstone.The Kunnam Member is arenaceous, composed of silty to sand claywith dispersed phosphatic nodules, interlayered mudstone, siltstoneand fine grained sandstone. The formation yields fossils of both marineinvertebrates and isolated vertebrates primarily sharks and marinereptiles (e.g., Underwood et al., 2011; Verma et al., 2012a). The occur-rences of ammonite index taxa such as Calycoceras newboldi in theOdiyam Member and Pseudaspidoceras footeanum and Eucalycoceraspentagonum in the Kunnam Member indicate a middle Cenomanianand the late Cenomanian to early Turonian ages, respectively(Ayyasami, 2006). However, on the basis of ammonites of theMammitesconciliatum and P. footeanum assemblage zones, a Cenomanian to earlyTuronian age was inferred for the Karai Formation by Phansalkar andKumar (1983). The following foraminiferal biostratigraphic zoneshave been recorded in chronological order within the formation:H. planispira of the early to middle Albian age, Rotalipora evoluta ofthe early to middle Cenomanian age, Rotalipora reicheli of the middleto late Cenomanian age, Whiteinella archaeocretacea of the lateCenomanian to early Turonian age and Marginotruncana helvetica toM. sigali of the early Turonian age (Narayanan, 1977; Tewari et al.,1996). Nagendra et al. (2002) documented planktic foraminifers suchas R. reicheli and Praeglobotruncana stephani from the marl beds ofthe Karai Formation and assigned a Cenomanian to middle Turonianage to this formation. On the basis of the oysters such asExogyra (Costagyra) costata, Pycnodonte vesiculosa and Rhynchostreonsuborbiculatum, the middle Cenomanian to early Turonian age wasalso proposed for this unit (Ayyasami, 2006). Recently, Bragina andBragin (2013) documented three radiolarian assemblages, namely,Halesium triacanthum to Orbiculiforma nevadaenis, Crucella latum toCryptamphorella micropora and Becus sp. B to Godia concava of the lateAlbian to middle Cenomanian age from the Karai Formation. The lime-stone deposition ceased suddenly in the late Albian due to an increasein basin depth as well as clastic input, and the presence of marine clayand silty clay having abundant gypsiferous clay and phosphatic nodulesalongwith amassive concentration of belemnites and ammonites in theKarai Formation is considered to be indicative of deposition that tookplace during increasing sea levels (Nagendra et al., 2002, 2011;Ramkumar et al., 2004). On the basis of foraminifers such as Astacolus,Gavelinella, Hoeglundina, Lenticulina and Nodosaria, Chidambaram(2000) proposed outer neritic to upper bathyal depositional conditionsfor the upper part of the formation. In addition, the occurrence of am-monites, belemnites and worm tubes indicate a deeper middle neriticenvironment (Nagendra et al., 2011). While proposing a revisedlithostratigraphic classification for the Cauvery Basin, Sundaram et al.



(2001) interpreted an offshore, highstand depositional environment.The analysis of oxygen isotope data of belemnites favours 16 to 18 °Cpalaeotemperature and some 150 to 200 m palaeodepth for the KaraiFormation (Zakharov et al., 2006). The presence of vertebrates suchas shark and marine reptile remains indicate a marginal marineenvironment. The coarser siliciclastic sediments on top of the Karai For-mation indicate that deposition occurred during a regressive phase(Ramkumar et al., 2004).

The Trichinopoly Group unconformably overlying the UttatturGroup is a discrete tectonostratigraphic unit, which has been separatedfrom the underlying Uttattur Group by a regional unconformity. It is upto 490 m thick and consists of conglomerates, shelly limestones, sand-stones and shales. The group is subdivided into the Kulakkalnattamand Anaipadi formations (Sundaram et al., 2001). The KulakkalnattamFormation represents a transgressive phase at the base of the groupand consists of sandstones, conglomerates, mudstones, shales andcoquinites. It is about 230 m thick and yielded invertebrates and drift-wood. The architecture of bioturbated, pebbly, cross-bedded sandstonesand shell-yielding conglomerates at the base of the formation thatin places contains drift-woods indicate a transgressive, littoral to shal-low marine environment. However, the occurrence of fine-grainedsiliciclastic sediments in the upper part of the formation shows shallowshelf deposition (Sundaram et al., 2001). The local abundance ofbivalves, gastropods, ammonites, fossil wood fragments and burrowssuggest that the unit was deposited under estuarine to intertidalcoastal conditions (Ramkumar et al., 2004; Nagendra et al., 2014). TheKulakkalnattam Formation is of Turonian age based on ammonites ofthe Romaniceras (Yubariceras) ornatissimum Zone and the oysterExogyra haliotoidea (Sundaram et al., 2001; Ayyasami, 2006). TheAnaipadi Formation unconformably overlying the Kulakkalnattam For-mation encompasses a regressive phase and consists of mudstones,sandstones and siltstone units. It is about 264 m thick, rich in molluscsparticularly in ammonites and rare marine reptiles. The formation ischaracterized by fine-grained and bioturbated siliciclastic sedimentswith abundant cephalopods, brachiopods and rare marine reptiles sug-gest deeper marine probably neritic conditions. The unit generallyshows a coarsening upwards trend, which reflects a large scale shoaling,regressive phase (Sundaram et al., 2001). The basal part of the unit hasyielded ammonites of the Lewesiceras anapadense and L. vaju zones andoyster Exogyra (Costagyra) fausta, and the upper part containsKossmaticeras theobaldianum, suggesting Turonian and Coniacian ages,respectively (Sastry et al., 1968; Ayyasami, 2006).

The Ariyalur Group rests unconformably on the Trichinopoly Groupand has its broadest distribution in the Cauvery Basin. It is subdividedinto three formations such as the Sillakkudi, Kallankurichchi andKallamedu in ascending order (Sundaram et al., 2001). The SillakkudiFormation dominantly consists of arkosic sandstones and limestonebands with abundant molluscs, indicating deposition during a trans-gressive phase in littoral to shallow marine conditions (Sundaramet al., 2001; Nagendra et al., 2014). This unit has yielded the ammonitessuch as Karapadites karapadense and the oyster Ostrea zitteliana, thebenthic foraminifers Bolivinoides culverensis and B. decoratus andthe planktic foraminifers Globotruncana elevata and Globotruncanaventricosa and is considered to be of Campanian age (Sastry et al.,1968; Govindan et al., 1996; Ayyasami, 2006). The contact betweenthe Sillakkudi and overlying Kallankurichchi formations is marked bythe presence of conglomerates, which may indicate a sea level fall atthe top of the Sillakkudi Formation. The Kallankurichchi Formation con-sists of conglomerates, limestones and sandstones, and rich in bryo-zoans, molluscs and foraminifers. The dominance of ferruginouslimestones and highly ferruginized foraminifers are indicative of a sealevel rise and deposition of the unit under oxidizing conditions(Nagendra et al., 2014). Calcareous sandstone bodies containing mol-lusc and bryozoan skeletons are interpreted to be deposited under shal-low marine conditions. The sediments of the upper part of the unit arerich in terrigenous input and in turn, indicate deposition during a

regressive phase (Sundaram et al., 2001). The occurrence of a dinosauregg in the shallow marine Kallankurichchi Formation is thought tohave been transported from some distance from its original placeinto the marine environment by streams or rivers based on the thick-ness of the eggshell (Kohring et al., 1996). The unit has yielded theammonite Hauericeras rembda, the oyster Pycnodonte (Phygraea)vesicularis and the planktic foraminifers Globotruncana gansseri andGlobotruncanastuarti, which are indicative for an early Maastrichtianage (Sastry et al., 1968; Govindan and Ravindran, 1996; Ayyasami,2006). The Kallamedu Formation comprises laminated sandstones,mudstones, red clays and sandy clays. The lower part of the unit yieldsabundant remains of molluscs, echinoids, bryozoans and planktic fora-minifers, indicating shallow marine depositional environments in aswallowing basin (Sundaram et al., 2001; Ramkumar et al., 2004). Theupper part contains palaeosols and abundant remains of the terrestrialand freshwater vertebrates such as frogs, turtles, crocodiles, dinosaursand a mammal, suggesting that the establishment of continental condi-tions and sediments appear to have been deposited under floodplainsettings (Ayyasami, 2006; Prasad et al., 2013). On the basis of the occur-rences of the ammonites Eubaculites vagina and Pachydiscus otacodensis,the oyster Agerostrea ungulata and nannofossils such as Arkhangelskiellacymbiformis and Ahmuellerella octoradiata, a late Maastrichtian age wasassigned to the lower part of the unit and a latestMaastrichtian agewasproposed for the upper vertebrate-yielding part of the unit, which isoverlain by the Niniyur Formation of the Danian age (Ayyasami, 2006;Prasad et al., 2013; Rai et al., 2013).

3. Cretaceous vertebrate diversity

Vertebrate remains have long been known from the Cretaceoussequences of the Cauvery Basin (e.g., Egerton, 1845; Blanford, 1862;Stoliczka, 1873; Lydekker, 1879), but only little consideration has beengiven to understand their diversity and palaeobiogeographic implica-tions to date. Recent reports of diverse fossil records from the CauveryBasin are remarkably improved the knowledge about vertebrate diver-sities and also added new important forms (Underwood et al., 2011;Verma et al., 2012a; Goswami et al., 2013; Prasad et al., 2013). In addi-tion, the new data also allows direct comparisons with contemporane-ous vertebrates known from other continents such as South America,Africa, Madagascar and Europe and thus leading to infer crucialpalaeobiogeographic patterns, in the context of the latest availablepalaeocoastline reconstructions of the Gondwanan continents. Thevertebrate fauna from theCauvery Basin is nowknown to includefishes,frogs, reptiles (ichthyosaurs, plesiosaurs, turtles and dinosaurs) and amammal (Table 2; Figs. 2–4).

3.1. Fishes

Fishes were the earliest discovered vertebrates from the CauveryBasin. During the investigation of Cretaceous fossiliferous sequences inthe Pondicherry and adjacent areas, south India between 1840 and1842, C.J. Kaye and C.E. Cunliffe collected fish remains from limestonebeds and handed over their collection to Sir Philip Egerton for systematicstudies (Blanford, 1862). Egerton (1845) identified twelve shark species,and one form of pycnodont and enchodont actinopterygian. Stoliczka(1873) documented the presence of durophagous shark, Ptychoduslatissimus Agassiz, 1843 and pycnodont,?Pycnodus sp., from the UttatturGroup of Ariyalur sub-basin, while Blanford (1862) also mentioned thepresence of Ptychodus and Pycnodus in the Uttattur Group. Later,Lydekker (1887) mentioned the occurrence of the same assemblage inCretaceous deposits of Ariyalur sub-basin and all these authors main-tained 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 taxonomicassignment and hence not mentioned. However, in most of the cases,

Table 2Vertebrate assemblage from the Cretaceous sequences of the Cauvery Basin.

Organism Taxon Locality Group Age References

Chondricthyes Sharks Pondicherry andadjacent areas

– ?Late Cretaceous Egerton (1845)

Uttattur Uttattur Albian to early Turonian Stoliczka (1873)Oxyrhina sp. Terani Uttattur Late Albian to early Turonian Paul (1973)Protosqualus sp., Gladioserratus magnus,?Notidanodon sp., Cretalamna appendiculata,Dwardius sudindicus, ?Eostriatolamin sp.,Squalicorax aff. baharijensis and Cretoduslongiplicatus.

Garudamangalam Uttattur Early Cenomanian Underwood et al. (2011)

Ptychodus latissimus Olapaudi Uttattur ? Late Cretaceous Stoliczka (1873)Ptychodus decurrens Garudamangalam Uttattur Late Cenomanian to early Turonian Verma et al. (2012a)

Osteichthyes Pycnodont and enchodont actinopterygian Pondicherry andadjacent areas

– ? Late Cretaceous Egerton (1845)

Lepisosteidae indet. Kallamedu Ariyalur Late Masstrichtian Prasad et al. (2013)Lycoclupea menakiaea Odiyam Uttattur Cenomanian Gowda (1967)Pycnodus sp. Uttattur Uttattur Late Albian to early Turonian Blanford (1862); Stoliczka (1873)

AmphibiansReptiles

Anura indet. Kallamedu Ariyalur Late Maastrichtian Prasad et al. (2013)Platypterygius indicusb Garudamangalam

and UttatturUttattur Late Albian to early Cenomanian Lydekker (1879);

Underwood et al. (2011)Dravidosaurus blanfordic Siranattam Trichinopoly Coniacian Yadagiri and Ayyasami (1979)Kurmademys kallamedensis Kallamedu Ariyalur Late Maastrichtian Gaffney et al. (2001)Testudines indet. Kunnam Uttattur Early Turonian Ayyasami and Das (1990)

Kallamedu Ariyalur Late Maastrichtian Prasad et al. (2013)cf. Simosuchus sp. and Crocodylia indet. Kallamedu Ariyalur Late Maastrichtian Prasad et al. (2013)Megalosaurus sp. Kallamedu Ariyalur Late Maastrichtian Lydekker (1879)Bruhatkayosaurus matleyib Kallamedu Ariyalur Late Maastrichtian Yadagiri and Ayyasami (1989)Megaloolithus cylindricusd Kallankurichchi Ariyalur Early Maastrichtian Kohring et al. (1996);

Fernández and Khosla (2014)Abelisauridae indet. and Troodontidae indet. Kallamedu Ariyalur Late Maastrichtian Prasad et al. (2013)

Mammal Sudamericidae indet. Kallamedu Ariyalur Late Maastrichtian Goswami et al. (2012)

a Otolith-based taxon.b Taxonomic validity is recently questioned.c Originally described as dinosaur taxon is now considered as a marine reptile.d Egg-based taxon.



precise locations and ages of fish-yielding horizons were not provided.Cretaceous rocks exposed at Pondicherry and Vridhachalam areaswhere Kaye and Cunliffe seemingly collected sharks teeth aremost likelychronologically aswell as lithologically to be coeval to the Ariyalur Groupof the Ariyalur sub-basin. The Cretaceous marine clastic rocks ofPondicherry and Vridhachalam regions are considered to be Santonianto Maastrichtian in age (Rasheed and Govindan, 1966; Banerji, 1968;Sundaram et al., 2001). It is, therefore, reasonable to assume that sharkremains described from the neighbourhood of Pondicherry area werecollected from the Santonian to Maastrichtian deposits. After a break ofnearly a hundred years, Paul (1973) described a shark tooth, Oxyrhinasp., from the Uttattur Group. More recently, Underwood et al. (2011)recorded some more marine shark species such as Protosqualussp., Gladioserratus magnus Underwood et al., 2011, ?Notidanodonsp., Cretalamna appendiculata Agassiz, 1843, Dwardius sudindicusUnderwood et al., 2011, ?Eostriatolamia sp., Squalicorax aff.baharijensis Stromer, 1927 and Cretodus longiplicatus Werner,1989 from the late Albian to Turonian Karai Formation (Fig. 3a–l).Additionally, Verma et al. (2012a) described one more species ofa durophagous shark, Ptychodus decurrens Agassiz, 1843, from theKarai Formation (Fig. 3m–n). Prasad et al. (2013) reported isolated gan-oid scales of lepisosteiformes from the freshwater deposits of theKallamedu Formation. Otolith-based fish species such as Lycoclupeamenakiae Gowda, 1967, belonging to actinopterygian and an un-named otolith as well as two ossiculiths described as specimens A andB were also reported from the Karai Formation (Gowda, 1964, 1966,1967).

3.2. Frogs

The record of amphibians from the Cauvery Basin is almostnon-existent as compared to frogs known from the Upper CretaceousDeccan volcanic province. Recently, a fragmentary ilium belonging to

an indeterminate anuran has been documented from the Maastrichtianfreshwater sediments of the Kallamedu Formation (Prasad et al., 2013).This is the only known report of any frog from the Cauvery Basin.

3.3. Reptiles

Marine reptiles are poorly known from the Cauvery Basin. Lydekker(1879) erected a new species of ichthyosaur, Platypterygius indicusLydekker, 1879, based on fifteen complete and fragmentary vertebraefrom the Karai Formation. Recently, Underwood et al. (2011) recovereda few isolated, complete teeth and partially fragmentary vertebrae,tentatively described as P. indicus (Fig. 3o–p). The authors, however,pointed out that a detailed examination is necessary to validate thetaxonomic status of this species. McGowan andMotani (2003) also pro-posed that material of P. indicus is not sufficient to erect a new speciesand considered the status of P. indicus as indeterminate. Yadagiri andAyyasami (1979) collected some skeletal material from Coniaciandeposits of the Anaipadi Formation, Trichinopoly Group on which theyerected a new stegosaur dinosaur, Dravidosaurus blanfordi Yadagiriand Ayyasami, 1979. Subsequent studies of the material doubted onits dinosaur affinity and suggested that these remains may belong to aplesiosaur (Chatterjee and Rudra, 1996; Galton and Upchurch, 2004;Paul, 2010; Wilson et al., 2011). Sahni (1957) described a fossilizedreptilian egg, referred to a chelonian that was recorded from theCenomanian sediments of the Uttattur Group.

The presence of fossil turtles in the Uttattur Groupwas first noted byMuzzy (1857) and Blanford (1862), but till date no turtle fossil has beenfully described from this group. However, a cast of the interior shell of aturtle was reported from the early Turonian sediments of the Karai For-mation (Ayyasami and Das, 1990). In discussing the cast of the turtle,the authorsmentioned that thematerial is not sufficient to be identifiedat species level, but observed that the cast of the animal had an elongat-ed shell, elevated in the vertebral region with sloping sides and had

Fig. 2. Generalized lithostratigraphic sections highlighting the probable stratigraphic positions of vertebrates, a. Karai Formation of the Uttattur Group (Lithocolumns modified afterBragina and Bragin, 2013) and b. Kallankurichchi and Kallamedu formations of the Ariyalur Group (Lithocolumns modified after Rai et al., 2013).



prominent sutural unions of the pelvis. Based on these observations, theauthors indentified it as a testudine pleurodiran turtle. In contrast, quiterecently two fossil turtles from theKallamedu Formationwere reported.Gaffney et al. (2001) described a single well-preserved, nearly completeskull and some postcranial remains of a turtle and assigned it to a newgenus and species Kurmademys kallamedensis Gaffney et al., 2001. It isa side-necked pelomedusid bothremydid and the authors also men-tioned that the material may include five other more poorly preservedand unprepared skulls. Subsequently, Gaffney et al. (2009) proposedthat three bothremydids such as Kurmademys from the Kallamedu,Sankuchemys from the Upper Cretaceous sediments associated withthe Deccan volcanism of Mumbai and Kinkonychelys from the UpperCretaceous Maevarano Formation of Madagascar are nested in thetribe Kurmademydini and are sister to each other. In addition, severalisolated, fragmentary carapaces and one vertebral piece recoveredfrom the Kallamedu Formation were referred to an indeterminatetestudine turtle (Prasad et al., 2013).

Despite more than 170 years of collecting vertebrates from theCauvery Basin, crocodylian remains were reported only recently fromthe Kallamedu Formation (Prasad et al., 2013). A single significant iso-lated multicuspid tooth referred to a Simosuchus-like notosuchian(Fig. 3q–r) and three isolated teeth alongwith one osteoderm identifiedas indeterminate Crocodylia have been described (Prasad et al., 2013).The report of a Simosuchus-like notosuchian from India is interestingas it represents the first occurrence of a Simosuchus-like crocodile out-side the Late Cretaceous of Madagascar. Simosuchus-like notosuchiansare only currently known from the Late Cretaceous of Madagascar andIndia (Prasad et al., 2013).

Dinosaurs are a significant biotic component of the Cauvery Basinand have been known since 1862, when Blanford discovered a singletooth and ill-preserved bones from the Kallamedu bone-bed of the

Kallamedu Formation (Blanford, 1862). Hundreds of dinosaur remainshave been collected from the Kallamedu bone-bed since then, thoughvery few of which have been identified and described. Lydekker(1879) identified the single isolated tooth of Blanford's collection as atheropod dinosaur and assigned it to the Megalosaurus sp. From thesame bone-bed, Matley (1929) reported large limb-bones and girdlesof sauropods, probably belonging to the Titanosaurus that was earlierrecorded from the Upper Cretaceous Lameta Formation of Jabalpur,Madhya Pradesh, central India and a few small bones assigned to stego-saurs. In addition, Matley (1929) also observed and mentioned thatbones occurring in the Kallamedu bone-bed were extremely friableand it was very hard to extract any complete, identifiable bones evenafter the greatest care was taken. Naryana Rao and Seshachar (1927)collected additional bones and concluded that these remains might be-long to dinosaurs. Later, Naryana Rao and Rama Rao (1930) jointly stud-ied a well-preserved vertebra from Naryana Rao and Seshachar'scollection and inferred that it represents the anterior part of a reptilianvertebra, probably belonging to a camarasaur sauropod. While re-examining the collection of Naryana Rao and Seshachar, Yadagiri et al.(1983) concluded that the collectionwas probablymammalian remainsfrom Pleistocene deposits; the occurrence of Pleistocene mammalianremains around the Kallamedubone-bedwas already earliermentionedby Matley (1929). An isolated well-preserved cervical vertebra recov-ered from theKallamedu Formationwas referred to theMegalosauridae(Rama Rao, 1932).

During the 1980s, geologists from the Geological Survey of Indiaexcavated the Kallamedu area and collected dinosaur material. Subse-quently, Yadagiri and Ayyasami (1989) described a new genus and spe-cies of a carnosaur theropod, Bruhatkayosaurus matleyi Yadagiri andAyyasami, 1989, based on material consisting of an ilia, an ischium, afemur, a tibia, a radius and a vertebra. The tibia is 2 m long and femur

Fig. 3. Selected Cretaceous vertebrates of the Cauvery Basin. a–n, shark taxa. a–d,Gladioserratus magnusUnderwood et al., 2011, isolated teeth. DUGF/2 (holotype) in labial (a) and lingual(b) views. DUGF/6 in labial (c) and lingual (d) views. e–h, Cretalamna appendiculata Agassiz, 1843, isolated teeth. DUGF/8 in labial (e) and lingual (f) views. DUGF/11 in lingual (g) andlabial (h) views. i–j, Dwardius sudindicus Underwood et al., 2011, isolated tooth, DUGF/21 (holotype) in labial (i) and lingual (j) views. k–l, Cretodus longiplicatusWerner, 1989, isolatedtooth, DUGF/40 in labial (k) and lingual (l) views. m–n, Ptychodus decurrens Agassiz, 1843, isolated teeth, PL/IGNOU/101 and 102 in occlusal views. o–p, Platypterygius indicus Lydekker,1879, isolated tooth, DUGF/41 in lingual (o) and labial (p) views. q–r, Notosuchid crocodile Simosuchus sp., isolated tooth, DUGF/48 in labial (q) and lingual (r) views. s–t, Troodontidaeindet., isolated tooth, DUGF/52 in labial (s) and lingual (t) views. u–v, Abelisauridae indet., isolated teeth, DUGF/53 in labial (u) and DUGF/54 in lingual (v) views (Figures a to l and o to pmodified after Underwood et al., 2011 and q to r and u to v after Prasad et al., 2013with permission from the Society of the Vertebrate Palaeontology,m to n after Verma et al., 2012awithpermission from Elsevier, and s to t after Goswami et al., 2013). DUGF stands for Delhi University, Geology Department, Fossil Catalogue, Delhi and PL/IGNOU for PalaeontologicalLaboratory, Indira Gandhi National Open University, New Delhi, India (where these fossil specimens are housed).



0.75 m, which are about 29% and 33% larger than the tibia and femur,respectively, of Argentinosaurus, a titanosaur sauropod from the LateCretaceous of Argentina, indicating that B. matleyi was larger thanArgentinosaurus and is one of the most gigantic dinosaurs ever discov-ered. However, some authors recently casted doubt on the taxonomicassignment of B. matleyi to a theropod dinosaur (Krause et al., 2006;Wilson et al., 2011). Krause et al. (2006) considered B. matleyi as asauropod dinosaur.

Kohring et al. (1996) described the earliest egg of an Indian dinosaurfrom the Kallankurichchi Formation. It is a reworked egg belonging tothe Megaloolithus cylindricus oospecies (Vianey-Liaud et al., 2003;Fernández and Khosla, 2014). Recently, Prasad et al. (2013) documentedfive isolated teeth of indeterminate Abelisauridae from the KallameduFormation (Fig. 3u–v). These teeth have a mesio-distally oriented long

axis, a strongly curved mesial profile and a straight distal profile anglingtoward the apex. Morphologically, the teeth are close to the ‘abelisauridMorphotype 3’ described from the Upper Cretaceous Maevarano Forma-tion of Madagascar (Fanti and Therrien, 2007; Prasad et al., 2013). Theinteresting and biogeographically significant discovery of troodontiddinosaur (Fig. 3s–t) from the Kallamedu Formation by Goswami et al.(2013) extends the record of troodontid dinosaurs from Laurasia to theLate Cretaceous of Gondwana, particularly in India.

3.4. Solitary mammal

A single tooth of a sudamericid gondwanatherianmammal has beenrecovered recently from the Kallamedu Formation (Goswami et al.,2012; Prasad, 2013). Gondwanatheria is an extinct group of non-

Fig. 4.Map showing the spatial distribution of the Albian to Maastrichtian vertebrate sites in India.



tribosphenic mammals that are reported only from the Gondwanancontinents, particularly South America, Antarctica, Africa, Madagascarand India (Verma et al., 2012b). Earlier, sudamericid mammals wereknown to occur in the Upper Cretaceous Deccan intertrappean beds ofMadhya Pradesh, Telangana and Karnataka (Prasad et al., 2007a;Verma et al., 2012b) and their presence in the Cauvery Basin representsthe southernmost extension of sudamericids in India (Fig. 4). However,other groups ofmammals such as adapisoriculids, archaic ungulates andharamiyids known from the Deccan volcanic province are yet to bedocumented from the Cauvery Basin (Prasad et al., 2006, 2007b;Khosla and Verma, 2014).

4. Palaeobiogeographic considerations

The Cretaceous sequences of the Cauvery Basin contain abiogeographically significant components of the vertebrate fauna ofIndia, adding to previously documented faunas from the Deccan volca-nic province (e.g., Khosla and Verma, 2014; Prasad and Sahni, 2014).An important result from the analysis of the Cretaceous vertebrates ofthe Cauvery Basin is that the latest Albian to Coniacian pre-Kallamedu(i.e., Uttattur and Trichinopoly) fauna is mostly marine in provenance,whereas the late Maastrichtian Kallamedu fauna is terrestrial tofreshwater. This has important bearings on understanding thepalaeobiogeographic role of the Indian plate during the Cretaceous.

Shark taxa such as Squalicorax (belonging to Family Anacoracidae),Dwardius and Cretalamna (Family Otodontidae), ?Eostriatolamia sp.(Family Odontaspididae), Protosqualus sp. (Family Squalidae),Gladioserratus and ?Notidanodon (Family Hexanchidae) andCretodus (Order Lamniformes, Family incertae sedis) are reportedfrom the Uttattur deposits. These were both active pelagic or ben-thic predators and usually inhabitants of tropical to temperate shallowwaters close to the shores, except Notidanodon, which was likely tolive in the deep waters of the continental shelf and slope (Cappetta,1987, 2012; Welton and Farish, 1993; Mannering and Hiller, 2008;Underwood et al., 2011). The anacoracid genus Squalicorax rangesfrom the Albian to Maastrichtian and is known from North America,South America, Europe, Russia, Asia, Africa, Australia and Madagascar(Welton and Farish, 1993; Cappetta, 2012; Figs. 5 and 6). Otodontidsharks Dwardius, was known from the Albian to the Late Cretaceous ofNorth America, Europe, Russia, Central Asia and Western Australia(Siverson, 1999; Underwood et al., 2011; Cappetta, 2012; Figs. 5 and6) and Cretalamna was reported from the Early Cretaceous to theEarly Eocene of Europe, North America, Asia, Russia, South America,Australia, Madagascar and Africa (Cappetta, 1987, 2012; Figs. 5 and 6).The odontaspidid Eostriatolamia was recorded from the Aptian to theLate Cretaceous of Europe, Central Asia and Australia (Cappetta, 1987,2012; Figs. 5 and 6). The squaliform shark, Protosqualus, has been re-corded from the Early to Late Cretaceous of Europe and Russia (Figs. 5and 6). The hexanchid shark, Notidanodon, was previously documented

Fig. 5. Stratigraphic distribution of Cretaceous sharks viz., Ptychodus decurrens, P. latissimus, Cretalamna, Squalicorax, Cretodus, Dwardius, Eostriatolamia, Protosqualus, Notidanodon and theichthyosaur Platypterygius. Mad — Madagascar and Ant — Antarctica.



from the Early to Late Cretaceous of Europe, North America, Africa, NewZealand, Australia and Antarctica (Underwood et al., 2011; Cappetta,1987, 2012; Figs. 5 and 6) and the other hexanchid shark,Gladioserratus,was reported from the Hauterivian to Cenomanian and possibly theEarly Eocene of Europe and Australia (Kemp, 1991; Underwood et al.,2011; Adolfssen and Ward, 2013). However, Cappetta (2012) offeredthe view that the Indian Gladioserratus is possibly a synonym of thegenus Notorynchus, which previously was reported from Cretaceousdeposits of Europe and Australia (Siverson, 2010). The lamniformCretodus, which probably disappeared in the Santonian, is well knownfrom the Albian to Santonian of Europe, North America, Central Asiaand Africa (Cappetta, 2012; Figs. 5 and 6). The durophagous shark,P. decurrens, was documented from the late Albian to Turonian depositsin North America, Europe, Africa, South America and Australia, whereasthe other species, P. latissimus, has been well known from the Turonian

to Campanian of North America, Europe, Russia and Asia (Niedžwiedzkiand Kalina, 2003; Cappetta, 2012; Verma et al., 2012a; Figs. 5 and 6).

The global occurrences at generic level of some shark taxa compris-ing Squalicorax, Notidanodon, Cretalamna, Cretodus and Ptychodus implythat these sharks had wide geographic distributions, inhabitanting di-verse climatic regions and were adapted to tropical and temperatecoastal life styles (Cappetta, 1987, 2012; Underwood et al., 2011). TheAlbian to Cenomanian record of some other shark taxa such asGladioserratus, Protosqualus, Dwardius, Notidanodon and Ptychodus aretypically high palaeolatitudes and found in northern Europe, northernNorth America, southern South America and Australia thus suggestingan ‘antitropical’ distribution in cooler waters (Underwood et al.,2011). Furthermore, their presence in southern India suggest thatthese sharks were frequent marine commuters in Cretaceous timesprobably from the latest Albian to early Turonian that roamed coastal

Fig. 6. Spatial distribution of Cretaceous sharks viz., Ptychodus decurrens, P. latissimus, Cretalamna, Squalicorax, Cretodus, Dwardius, Eostriatolamia, Protosqualus, Notidanodon and theichthyosaur Platypterygius shown on a Late Cretaceous (94 Ma) palaeogeographic map (map modified from Scotese, 2001).



regions of the Tethyan Sea as well as the Indian Ocean. Recent reportsabout the equatorial occurrence of Ptychodus decurrens and Cretalamnasp. from the Late Cretaceous (early Turonian and ?Coniacian toSantonian) of Nigeria reflect the occurrence of episodic cooler waterevents that may have facilitated their dispersals between high-palaeolatitudes of both hemispheres (Vullo and Courville, 2014).

The marine reptile, Platypterygius, is one of the most geographicallywidely distributed and stratigraphically long-lived Cretaceous ichthyo-saur genera, as it has been reported from the Hauterivian to Barremianof South America, Aptian to Albian of Australia and Albian to Cenomanianof Europe, Russia and North America (e.g., Zammit, 2012; Figs. 5 and 6).The latest Albian to Turonian occurrence of Platypterygius in Indiaindicates a cosmopolitan distribution and wide marine territory. Whilecomparing the Cretaceous megainvertebrate fauna of the CauveryBasin with those known from Europe, the Mediterranean region andAfrica, Kossmat (1895) also visualized a similar distribution pattern forinvertebrates.

The latest Albian to Coniacian marine vertebrate fauna of theCauvery Basin has intracontinental affinities with those occurring inthe Bagh Formation, Narmada Basin, central western India (Verma,1965). The recovered faunaof theBagh Formation comprises indetermi-nate lamniform and ptychodont (“Ptychodus mahakalensis”) sharks andpycnodont (“Coelodus malwaensis”) fishes (Verma, 1965, 1968, 1969;Dassarma and Sinha, 1966; Chiplonkar and Ghare, 1977). The BaghFormation is a shallow marine Upper Cretaceous sequence that wasdeposited in an east–west direction, extending from Madhya Pradeshto Gujarat in the Narmada Basin. Stratigraphically, it is divided intothe Nimar Sandstone, Nodular Limestone, Deola-Chirakham Marl andCoralline Limestone in ascending order. Interestingly, almost all fisheswere recovered from the oyster bed that lies between the Nimar Sand-stone and Nodular Limestone, exposed near Amba Dongar and theKawant areas in Gujarat and the Mahakal area in Madhya Pradesh(Verma, 1965; Dassarma and Sinha, 1966; Chiplonkar and Ghare,1977; Fig. 4). The contact between the Nimar Sandstone and the overly-ing Nodular Limestone is conformable and has yielded many inclusionsof a marine transgression event (Jafar, 1982; Bose and Das, 1986; Taylorand Badve, 1995). Chiplonkar et al. (1977) assigned an Albian toCenomanian age to the Nimar Sandstone and the Coniacian to Turonianto the Nodular Limestone based on placenteritid ammonoids. On the

basis of nanoplanktons, Jafar (1982) suggested that the complete BaghFormation was deposited during the late Turonian transgression. Alate Turonian to Coniacian age has also been assigned to the top oysterbed based on the occurrence of the ammonite, Barroisiceras onilahyense(Gangopadhyay and Bardhan, 2000). Khosla et al. (2003) assigned aCenomanian to Turonian age based on the presence of fragmentaryskeletal remains of sauropod dinosaurs from the Bagh area, DistrictDhar, Madhya Pradesh. More recent data favours a Cenomanian agefor the Nimar Sandstone and Turonian age for the overlying NodularLimestone (Jaitley and Ajane, 2013). In addition to sharks, the marineplesiosaur reptile, Thaumatosaurus indicus, has been recovered fromthe Umia Formation exposed at Umia (Fig. 4) in the Kachchh Basin,Gujarat (Lydekker, 1889; Verma et al., 1977). A shallowmarine deposi-tional environment was inferred for the Umia Formation and it wasdated to be of late Tithonian to Albian age (Jaikrishna et al., 1983).

The Bagh and Umia formations of the western margin of Indiacontain some marine vertebrates which are closely related to thosefrom the probably coeval sediments of the Cauvery Basin, southeasternIndia and thus suggest some latest Albian to Coniacian faunal inter-dispersal events between these basins. Numerous studies show thatthe sea level was very high during the latest Albian to Campanian(e.g., Miller et al., 2005), flooding the continents. India had alsowitnessed two important Albian to Campanian transgression events;one in the Narmada valley that came from the west and resulted inthe deposition of the Bagh and Umia formations and second on theeast coast of India that formed the Cauvery Basin and probably persistedto the beginning of the late Maastrichtian (Acharyya and Lahiri, 1991).The late Albian to Turonian invertebrate faunas including nautiloids,ammonoids, bivalves, gastropods and echinoids of the Cauvery andNarmada basins are not only closely related to each other, but alsobear similarities with those known from the contemporaneous depositsofMadagascar, North Africa and Europe (Chiplonkar and Tapaswi, 1979;Smith, 2010). Here, it is proposed that intermingling between theCauvery, Bagh and Kachchh marine faunas may have taken place bytwo possible sea routes: first via the Mozambique Channel, linking thesouthern Tethyan Sea to the northern Indian Ocean and second, via aninitial shallow water connection that may had been establishedbetween southern Tethyan Sea and northern Indian Ocean along thecoast of Cape Comorin and Sri Lanka via the Mascarene Basin prior to

Fig. 7. Stratigraphic distribution of Cretaceous vertebrates viz., kurmademydine turtles,Simosuchus-like notosuchian crocodiles, troodontid and abelisaurid dinosaurs andsudamericid mammals. SA — South America, Mad — Madagascar, Eu — Europe andEu + As + NA — Europe, Asia and North America.



the complete separation of the Indo-Madagascar block, possibly relatedto initial rifting and rise in sea level (e.g., Smith et al., 1994; Scotese,2004; Fig. 6). These sea routes would have facilitated the inter-dispersal of marine faunas and rise of sea level along with localsubsidence that may have displaced them into these basins. In contrast,the presence of pycnodont fish in the Uttattur Group and the Bagh For-mation extends its record to the latest Albian to Coniacian times in India

Fig. 8. Spatial distribution of Cretaceous vertebrates viz., kurmademydine turtles, Simosuchus-likshown on a latest Cretaceous (65 Ma) palaeogeographic map (Map modified from Blakey, 200

(Stoliczka, 1873; Chiplonkar and Ghare, 1977). Earlier, pycnodont fos-sils broadly documented from the Maastrichtian sites such asDongargaon, Nagpur (Maharashtra); Rajahmundry (Andhra Pradesh);Marepalli, Timsanpalli (Telangana); Kisalpuri, Lotkheri (MadhyaPradesh); Kora (Gujarat) and Papro (Uttar Pradesh) of the Deccan vol-canic province (Fig. 4) were considered to be the oldest records fromIndia and their presence in the Cauvery Basin was not taken intoaccount (see Prasad, 2012). It is possible that Maastrichtian pycnodontsof the Deccan volcanic province were the probable immigrants of theCauvery and Bagh pycnodonts that were already present in coastalregions of the continent. In most of the sites, pycnodonts were foundin association with the myliobatid ray, Igdabatis, in the Deccan volcanicprovince and both fish are considered inhabitants of pelagic, coastalmarine and shelf environments (e.g., Welton and Farish, 1993; Prasad,2012; Martin-Abad and Poyato-Ariza, 2013). However, freshwaterpycnodonts were also documented in the fossil record (Prasad, 2012;Martin-Abad and Poyato-Ariza, 2013). The presence of Igdabatis andpycnodonts in the Deccan volcanic province points to the fact thatshallow marine incursions may have occurred in the these ray-yielding sites possibly through the Narmada–Tapti and Godavari rifts,and suchmarine incursionswould have dispersed pycnodonts into con-tinental areas during the Maastrichtian (sensu Sahni, 1983). Recently,similar marine incursions at the Cretaceous/Palaeocene transitionwere documented based on foraminifers recorded from the Jhilmili(Madhya Pradesh) and Rajahmundry (Andhra Pradesh) sites of theDeccan volcanic province and the Krishna-Godavari Basin, respectively(Keller et al., 2009).

Three Gondwanan groups: bothremydid turtles, notosuchian croco-diles and sudamericidmammals, are particularly important elements ofthe non-marine Kallamedu fauna as they provide stronger evidence tosupport the similarities between Indian and Madagascan faunas, andto some extent with South American (Gaffney et al., 2009; Prasadet al., 2013; Figs. 7 and 8). The bothremydid turtle, K. kallamedensis, isconsidered to be a sister-taxon of Kinkonychelys rogersi from the LateCretaceous of Madagascar and Sankuchemys sethnai from the LateCretaceous of India (Gaffney et al., 2009). These three taxa placedtogether in a single tribe, Kurmademydini, have only been reportedfrom the Late Cretaceous of Indo-Madagascar. In addition, the discovery

e notosuchian crocodiles, troodontid and abelisaurid dinosaurs and sudamericidmammals6).



of a Simosuchus-like notosuchian crocodile from the Kallameduadds new evidence in favour of bioconnections between India andMadagascar in the latest Cretaceous as Simosuchus has only previouslybeen reported from the Late Cretaceous of Madagascar (Buckley et al.,2000; Prasad et al., 2013; Fig. 8). The presence of sudamericidgondwanatherian mammal in southern India not only strengthens thesimilarities of faunas between Indo-Madagascar and South America,but also broadens the geographic distribution of the group withinIndia (Prasad et al, 2007a; Verma et al., 2012b; Fig. 4). The report of aLaurasian clade including maniraptoran troodontid dinosaur from theKallamedu is themost intriguing as it represents the first Gondwana re-cord of troodontid dinosaurs that extends the geographic range of thegroup into the Late Cretaceous of India from Laurasia (Goswami et al.,2013; Figs. 7 and 8). Further, it raises several issues such as whethertroodontids dispersed to India from Laurasia during the Late Cretaceousor whether the group had an unknown broader Gondwanan distribution.Goswami et al. (2013) argued that since the oldest record of troodontidsis coming from the Middle Jurassic of Laurasia, their presence in the LateCretaceous of Indiawould have been a result of vicariance or dispersal as-sociated with the movement of the tectonic plates.

The occurrence of abelisaurid dinosaurs from the Kallamedu is quiteinteresting as it represents the southern extension of the Indianabelisaurids (Fig. 4). Indian abelisaurids, Indosuchus raptorius andRajasaurus narmadensis, known from the Upper Cretaceous Lameta For-mation of Madhya Pradesh and Gujarat (Fig. 4), respectively, havesister-group relationships with Majungasaurus crenatissimus, anabelisaurid from the Upper Cretaceous Maevarano Formation ofMadagascar (Wilson et al., 2003). In addition, Indo-Madagascanabelisaurids are closely related to South American taxa thus suggestinga Late Cretaceous biogeographic connections between them (Carranoand Sampson, 2008). Other than the Kallamedu fauna, the presence ofabelisaurid dinosaurs, leptodactylid and ranoid frogs, pelomedusoidturtles and madtosiid, and nigerophiid snakes in the Upper Cretaceoussediments associatedwith theDeccan volcanismhavingGondwanan af-finities particularly with South American and Madagascan groups,strongly indicates cosmopolitanism among these Gondwana faunas(Krause et al., 1997; Prasad, 2012; Khosla and Verma, 2014). Prior to2014, unquestionable remains of abelisaurid dinosaurs were knownonly from Gondwanan continents, but recent reports of abelisauridtaxa, Arcovenator escotae Tortosa et al., 2014, from the Late Cretaceousof France (Fig. 8), may weaken the concept of Gondwana endemismfor abelisaurid dinosaurs (Tortosa et al., 2014). Moreover, A. escotae iscloser to the Indian and Madagascan abelisaurids than to SouthAmerican taxa. Tortosa et al. (2014) maintained that the presence ofan abelisaurid in the Late Cretaceous of Europe possibly reflects two sce-narios: either abelisaurids had a Pangaean distribution or it dispersedfrom Africa to Europe in the beginning of the Late Cretaceous as therewere frequent biotic dispersal routes between these continents(e.g., Gheerbrant and Rage, 2006). The discovery of a new abelisauridsuch as Eoabelisaurus mefi Pol and Rauhut, 2012 from the Middle Juras-sic of Patagonia extended the record of the group back by more than40 Ma in Gondwana (Pol and Rauhut, 2012). Nevertheless, the discov-ery of a troodontid from the Late Cretaceousmay be explained by a sim-ilar dispersal model as proposed for A. escotae. A recentpalaeogeographic reconstruction proposes that India was connected toEast Africa through theOman-Kohistan-Dras Island Arc in the latest Cre-taceous (Chatterjee et al., 2013) that would have provided a dispersalcorridor for biotic exchanges between India and Africa. In addition, thepresence of definite and undisputed Gondwanan adapisoriculid euthe-rians such as Deccanolestes in the Late Cretaceous of India having closerelationships with Afrodon (Adapisoriculidae) from the Late Palaeoceneof Africa and Europe suggests the latest Cretaceous faunal dispersals be-tween India and Africa (Khosla et al., 2004; Verma, 2008; Boyer et al.,2010; Prasad et al., 2010; Goswami et al., 2011). Finally, it is prematureto conclude, whether dispersal or vicariance events played a role inshaping the Late Cretaceous vertebrate biogeography of India in

particular and Gondwana in general because the Lower Cretaceous sed-iments of Indo-Madagascar and South America, and the Upper Creta-ceous sediments of Africa and Australia are not well-sampled.

5. Conclusions

An overview of the vertebrates from Cretaceous (Albian toMaastrichtian) successions of the Cauvery Basin, Tamil Nadu, southernIndia shows that the Uttattur and Trichinopoly groups, ranging fromthe Albian to Coniacian, yielded mainly marine vertebrates and theMaastrichtian Kallamedu Formation of the Ariyalur Group is known toyield non-marine vertebrates. The marine vertebrate assemblagesconsist of sharks (Squalicorax, Dwardius, Cretalamna, ?Eostriatolamia,Protosqualus, Gladioserratus, ?Notidanodon, Cretodus and Ptychodus),pycnodonts (Pycnodus), enchodonts, otolith-based actinopterygian(Lycoclupea menakiae), ichthyosaurs (Platypterygius indicus), plesio-saurs (Dravidosaurus blanfordi), testudine turtles and a reptilian eggshell. Many taxa show cosmopolitan distributions having broaderbiogeographical affinities to Laurasian continents, and to some extentwith Gondwana. The reports of pycnodont fishes (Pycnodus and“Coelodus”) from the shallow marine Uttattur Group and the Bagh For-mation extends the Indian record of pycnodonts from the lateMaastrichtian to the latest Albian and Coniacian and it might havebeen possible that Maastrichtian pycnodonts from the Deccan volcanicprovince were the immigrants of pycnodonts of the Cauvery Basin andthe Bagh Formation. The non-marine vertebrate assemblage comprisesfrogs, turtles (Kurmademys kallamedensis), crocodiles (Simosuchus-likenotosuchian, Crocodylia indet.), dinosaurs (Megalosaurus sp., Titanosaurussp., B. matleyi, Megaloolithus, Abelisauridae indet. and Troodontidaeindet.) and a mammal (Sudamericidae indet.). The Kurmademys turtleand Simosuchus-like crocodile are suggestive of close biogeographicconnections between India andMadagascar during the latest Cretaceous.The abelisaurid dinosaurs and sudamericid gondwanatherian mammalsupport faunal similarities during the Late Cretaceous between SouthAmerica and Indo-Madagascar. The presence of a troodontid dinosaur inthe Late Cretaceous of India and an abelisaurid dinosaur in Francemay in-dicate sporadic dispersals between India and Europe via Africa. Thoughabelisaurid and troodontid dinosaurswere present in the Late Cretaceousof France and India, respectively, the abelisaurids were clearly the moredominant components of Gondwanan continents particularly, SouthAmerica, Africa, Madagascar and India whereas the troodontids werethe more dominant faunal elements of Laurasian continents such asAsia, Europe and North America.

Acknowledgements

The author expresses his sincere thanks to Ashok Sahni(Lucknow), G.V.R. Prasad (Delhi), Ashu Khosla (Chandigarh), R.Nagendra (Chennai), J. Pollerspöck (Germany) and S. Hamm(Texas) for their advice and fruitful discussions that have consider-ably improved the manuscript. Special thanks go to Avtar Changotra(Mumbai), K.G. Kulkarni (Pune), R. Nagendra (Chennai), A.S. Rathore(Delhi), J. Rai (Lucknow), M. Ramkumar (Salem) and J. Pollerspöck(Germany) for providing useful literature. Vijayshri and AseemKumar (Delhi) are gratefully acknowledged for providing necessaryfacilities and supports. Author is grateful to S.K. Shah, V. Parmar(Jammu), J.J.W. Sertich (Denver) and C.J. Underwood (London) forlinguistic improvements of the manuscript. K. Sivakumar, T.Ayyamperumal (Chennai) and K. Singh (Jammu) are thanked fortheir assistance in field. Mansi Bangia is thanked for drafting somefigures. This research was supported by funds from the Departmentof Science and Technology, Government of India, Grant No. SR/FTP/ES-33/2008 (Fast-Track Research Project). The author wishes tothank the editor and the two anonymous reviewers for their con-structive comments.



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