GAIA N' 15, LlSBOAlLISBON, DEZEMBRO/DECEMBER 1998, pp. 379-387 (ISSN: 0871-5424)

STATUS AND PALAEOBIOLOGY OF THE LATE INDIAN THEROPODS WITH DESCRIPTION OF A NEW THEROPOD EGGSHELL OOGENUS AND OOSPECIES, ELLIPSOOLITHUS KHEDAENSIS, FROM THE , DISTRICT KHEDA, , WESTERN

Raminder S. LOYAL Centre of Advanced Study in Geology, Panjab University. CHANDIGARH· 160014. INDIA

Dhananjay M. MOHABEY Palaeontology Division, Geological Survey of India, Seminary Hills. NAGPUR - 440 006. INDIA

Ashu KHOSLA Centre of Advanced Study in Geology, Panjab University. CHANDIGARH - 160014. INDIA

Ashok SAHNI Centre of Advanced Study in Geology, Panjab University. CHANDIGARH -160014. INDIA

ABSTRACT: This paper highlights the Cretaceous theropod record in India along with its af­ finities, and palaeobiologic aspects including description of a new theropod eggshell from the Upper Cretaceous Lameta Formation of Gujarat, India. Recent discoveries and phyloge­ netic reassessment of various theropod families suggest the presence of the poorly known family ; recent reports confirm the presence of crenatissimus (DEPRET, 1896), HUENE & MATLEY, 1933 possible BONAPARTE, 1985 and ? MARSH, 1877. An updated list of various Cretaceous theropod genera in­ cludes abelisaurids (Indosuchus raptorius HUENE & MATLEY, 1932, Indosaurus matleyi HUENE & MATLEY, 1933, Majungasaurus crenatissimus (DEPRET, 1896) Carnotaurus sp., allo­ saurids (Compsosuchus solus HUENE, 1932, ?Allosaurus) and coelurosaurs (Jubbulpuria tenuis HUENE & MATLEY, 1933, indicus HUENE & MATLEY, 1933). Other theropods, such as Ornithomimoides mobilis HUENE & MATLEY, 1933, O. (?)barasim/ensis HUENE & MAT­ LEY, 1933 and Dryptosauroidesgrandis HUENE, 1932 regarded as indeterminate, need further assessment for proper affiliation. In the ongoing discussion, we however, present signifi­ cant new data on the first ever report of eggshell material referable to a new ooge­ nus and oospecies Ellipsoolithus khedaensis, (oofamily ) from near Rahioli, district Kheda, Gujarat. This material has been presently assigned to the and can be easily differentiated from the other well known megaloolithid oospecies (broadly referable to due to associated titanosaurid bones; known from Madhya Pradesh, Gujarat, Maharashtra and Ariyalur in South India) on the basis of megascopic, microscopic and ultrastructural characteristics. Megascopically, the eggshell shape varies from elon­ gated ellipsoid, having an elongation ratio range of 1.5 to 1.7. The eggshell thickness is 1.2- 1.6 mm and the mammillary layer is 1/4-1/7 of the total thickness. At the microstructural level, an angusticanaliculate pore system is observed. As the present find constitutes the first record of elongatoolithid eggshells apart from well known Chinese and Mongolian lo­ calities, comments on possible sauropod-theropod palaeobiology are also discussed.

379 artigos/papers R.S. LOYAL; O.M. MOHABEY; A. KHOSLA & A. SAHNI

THEROPOD RECORD IN INDIA AND gascar) including cross-sectional and serrational EGGSHELL STATUS morphologies are strongly similar to that of the In­ dian theropod Indosuchus raptorius HUENE & MAT­ Although Indian sauropod eggshell lEY, 1933; for instance, both forms exhibit marked and skeletal discoveries have made considerable pattern of vertical ridges on the interdental plates. As advancements overthe lastfew decades, the thero­ such, the shared synapomorphies between these pods unfortunately have received scant attention theropods have led SAMPSON et al. (1996) to consid­ except for notable exceptions (HUENE & MATlEY, erthem as sister taxa within the Abelisauridae. 1933; CHATTERJEE, 1978; CHATTERJEE & RUDRA, This viewpoint is also corroborated by CHATTERJEE 1996). Except for the presence of a small & RUDRA (1996); in a detailed discourse on Gon­ theropod Alwalkeria maleriensis CHATTERJEE & dwana theropod relationships, these workers asso­ CREISlER, 1994 from the Maleri Formation ciated a more primitive phylogenetic position to the (Pranhita-Godawari Valley, district Adilabad, abelisaurids in comparison to the allosaurs and ty­ Andhra Pradesh), the dominant discoveries of rannosaurs. Further, the remains of Majungasaurus theropod skeletal remains are associated with the crenatissimus (Mahajanga Basin, Upper Creta­ Upper Cretaceous Lameta and Intertrappean se­ ceous) represented by one premaxilla has been quences outcropping in central and southern parts confirmed to belong to an abelisaurid with no "mega­ of peninsular India. Although no theropod losaurian" ortyrannosaurid characteristics (KRAUSE record exists, the later theropods are stratigraphi­ & DODSON , 1994; KRAUSE & HARTMAN, 1996). More cally associated with the Upper Cretaceous Lameta recently, CHATTERJEE & RUDRA (1996: 517) believe and Intertrappean sequences, outcropping in cen­ that "Indosuchus is not a tyrannosaurid as was sup­ tral and southern parts of Peninsular India. We pro­ posed earlier, but possibly an abelisaurid". These vide below a brief overview of theropod groups workers discovered a complete skeleton of Indo­ prevailing in the Indian subcontinent during the Cre­ suchus from near Rahioli, Gujarat and referred it to taceous times. allosaurids on the basis of similar lengths of fore­ The Indian Cretaceous theropods are broadly limbs in these two taxa; however, hindlimb elements represented by skeletal remains of abelisaurids, al­ resemble those of Camotaurus, a well-established losaurids, coelurosaurs and some indeterminate abelisaurid. forms (HUENE & MATlEY, 1933; BUFFETAUT, 1987; Further, the other theropod known from Bara KRAUSE & HARTMAN, 1996; CHATTERJEE & RUDRA, Simla Hill, Indosaurus mat/eyi, bears thickened fron­ 1996). Amongst these forms, the taxonomic affini­ tals and elevated parietals; these features are also ties of larger theropods have remained in doubt, observed in Camotaurus sastrei, a well established awaiting further assessment; in particular "megalo­ and a diagnostic abelisaurid known from Goro Frigio saurids" from Kallamedu, south India (BLANFORD, Formation (Albanian-Cenomanian) of Chubut, Ar­ 1862; LYDEKKER , 1879), Takli (Maharashtra, gentina (BONAPARTE, 1985). Moreover, vertebral VIANEY-LiAUD, JAIN & SAHNI, 1987) and Rahioli elements closely similar to that of Camotaurus are (MATHUR & SRIVASTAVA, 1987). However, in com­ also known from Rahioli, Gujarat (CHATTERJEE & parison, the family Abelisauridae is a relatively bet­ RUDRA, 1996). ter known group from the Carnosaur Bed of Lameta Formation, Jabalpur and is represented by Indo­ The remaining forms from Bara Simla Hill locality suchus raptorius HUENE & MATlEY, 1933, Indosau­ by HUENE & MATlEY (1933)were referred to coeluro­ rus mat/eyi HUENE & MAllEY, 1933 and saurs, which included Compsosuchus solus HUENE, Camotasaurus BONAPARTE , 1985. Of these, the first 1932, Laevisuchus indicus HUENE & MATlEY, 1933, two forms, Indosuchus HUENE & MATlEY, 1933 and Jubbulpuria tenuis HUENE & MATlEY, 1933, Coelu­ Indosaurus HUENE & MATlEY, 1933 originally re­ roides largus HUENE, 1932, Oryptosauroides gran­ ferred to (HUENE & MATlEY, 1933), dis HUENE, 1932, Omithomimoides mobilis HUENE & have been variously assigned, for instance Indo­ MATlEY, 1933 and O. (?) barasimlensis HUENE & suchus as a tyrannosaur and Indosaurus as a mega­ MATlEY, 1933. Of these forms, Compsosuchus so­ losaur (CHATTERJEE, 1978; BUFFETAUT, 1987); and Ius was suggested by MOLNAR, KURZANOV & DONG more recently to abelisaurids (BONAPARTE & NOVAS, (1990) and MOLNAR & FARLOW (1990) to be an allo­ 1985; MOLNAR, 1990; MOLNAR, KURZANOV & DONG , saurid and particularly close to Allosaurus fragilis, 1990 and SAMPSON et al., 1996). on the basis of shared apomorphies. Recently, however, morphologic and phyloge­ From Bara Simla Hill locality, the only forms to netic comparison of various theropods including have retained their assignment to coelurosaurs are abelisaurids has been attempted by SAMPSON et al. Jubbulpuria tenuis and Laevisuchus indicus, but (1996). In this work, it has been considered that pre­ thought to be indeterminate (NORMAN, 1990). Other maxillary dentition of Majungasaurus crenatissimus forms which remain unidentified and await assess­ (DEPRET, 1896) (from Upper Cretaceous of Mada- ment are Coeluroides largus, Oryptosauroides

380 STA TUS AND PALAEOBIOLOGY OF THE INDIAN THEROPODS

2f s'

.Lavario ~ Muwodo

, ,~ Ohl011.-

, . timnton. (Colcrete ) 23 f~~~~ ~ _H ..U :!w :-:...... :-:-:.: . a-ritty Sand~tont ~ 0=0_' , , km 1_0~00 • • • • • • Oronite r • • •

~ Lom.to Formation (LotI' Cr.tQcI'OU~ )

~ Nf'!tho n ••ly found I'llfp!toid 1'99\.

I~I Ntnt of earlier known 1ophl'riCo' dino\our 1'99", I:::J ISOnH of Titano!tDurid" and The-ropod"

Fig. 1 - Location of newly found ellipsoidal eggs (elongatoolithid) in Kheda district, Gujarat. grandis, Omithomimoides mobilis and O. (?) bara­ more recently elongatoolithid and megaloolithid egg simlensis. Further, D. grandis is considered by MOL­ nests have been recorded from the same stra­ NAR (1990) to possess no carnosaurian characters, tigraphic level in this area. At least 60 eggs (ellipsoi­ while ornithomimosaurids are not present south of dal) have been located in more than half a dozen Mongolian localities (BARSBOLD & OSMOLSKA, nests in addition to stray eggs and eggshell debris 1990). (MOHABEY, 1996b.) A single nest has yielded thir­ teen eggs. In a single nest the eggs are disposed PRESENT EGGSHELLS AND NESTS: horizontally in a single layer (Fig. 2C). LOCALITY, STRATIGRAPHY AND FIELD DISPOSITION SYSTEMATICS Recent palaeontological work near Rahioli (Dis­ trict Kheda, Gujarat) led to acquisition of a large number of ellipsoid eggs both in isolated form as well Oofamily as in nests (Fig. 2-3). The present eggshell locality is Elongatoolithidae ZHAO, 1975 situated west of Lavaria Muwada and south of Keva­ Oogenus diya at a distance of nearly 1.5 km NW of Rahioli (Fig. 1). Ellipsoolithus n. oogen. The egg-bearing horizon is confined to the up­ permost calcretized sandstone. Stratigraphically, Diagnosis: Ornithoid basic ; ratite morpho­ this bed lies 1 m below the level, that had earlier type; angusticanaliculate pore system; outer sculp­ yielded spherical-shaped dinosaur eggshells be­ ture is lineartuberculate and dispersituberculate; longing to the oofamily . However ellipsoid eggs (elongation ratio 2) varying in diame-

381 R.S. LOYAL; D.M. MOHABEY; A. KHOSLA & A. SAHNI

ter from 98-110 mm and 65-80 mm at polar and thick wedges in the mammillary layer. The mammil­ equatorial region respectively; eggshell thickness lary layer is about 1/6-1/7 but more commonly 1/4 of 1.2-1.6 mm; two-layered eggshell; mammillary layer the total shell thickness. The spheroliths are broad comprises 1/4 -1 /7 of the total eggshell thickness. having wide domal - shaped roofs. The spheroliths are fused. and separated in the mammillary zone. Etymology: The oogenus Ellipsoolithus is Under SEM, thick wedge-shaped crystallites are named after the predominant ellipsoidal shape of seen in the mammillary layer. The upper spongy the eggs. layer shows gently curving growth lines towards the upper part olthe spheroliths. The pore system is an­ Ellipsoolithus khedaensis n. OOSp. gusticanaliculate. (Fig . 5 A-E; Fig . 6) Remarks: The eggshells and nests assigned to Holotype: An almost complete egg (Reg. No. the oofamily Elongatoolithidae are the most domi- 1491/CRP/96, housed at GSI, Nagpur). Etymology: The name khedaensis is derived af­ ter District Kheda, Gujarat. A Type locality: West of Lavaria Muwada and south of Kevadiya and at a distance of 1.5 km NW of Rahioli, district Kheda, Gujarat. Type horizon: Lameta Formation; Upper Sandy Carbonate (calcretized paleosol); Upper Creta­ ceous. Referred material: Many near complete to par­ tial eggs and eggshell debris. Repository: More than a dozen more or less complete and partial eggs are housed at Geological Survey of India, Nagpur with Dr. D. M. Mohabey; a few almost complete eggs are with Dr. Raminder S. Loyal, Centre of Advanced Study in Geology, Pan­ B jab University, Chandigarh. Diagnosis: Ellipsoidal eggs assuming near oval - shape, polar and equatorial diameter range 98-11 0 mm and 65-80 mm; elongation ratio ranging be­ tween 1.5-1.7 mm; thickness 1.2-1.6 mm; ornamen­ tation variab le along egg surface, disper­ situberculate to ramotuberculate at the polar region, lineartuberculate on the equatorial portion; two­ layered; mammillary layer ranges 1/4-1/7 of total eggshell thickness but more commonly 1/4; bulbous mammillae; spheroliths with curving domes. Description: Eggs are predominantly ellipsoidal c in shape and assuming a near oval-shape. Polar and equatorial diameters ranges between 98-110 mm , and 65-80 mm respectively (Fig. 4). The eggshell , thickness varies from 1.2-1.6 mm. Surface orna­ mentation comprises dispersituberculate to ramotu­ berculate patterns at the polar region, depicting \~ irregularly dispersed and ramified ridges and nodes. At the equatorial region, lineartuberculate ornamen­ tation is observed, with linear (and sometimes dis­ continuous) arrangement of tubercular nodes and Fig. 2 - Egg nests from near Lavaria Muwada locality, lineations, which are elongated parallel to the longer district Kheda, Gujarat; Lameta Formation; Upper Sandy axis of the eggshell (MOHABEY, 1996b). Carbonate (calcretized paleosol); Upper Cretaceous. A- B - Disposition of elongatoolithid eggs in different nests. The microstructure of the eggshell is double­ C - Section of three horizontally embedded eggs in the layered with radiating structures and presence of sandy carbonate.

382 STA TUS AND PALAEOBIOLOGY OF THE LATE CRETACEOUS INDIAN THEROPODS

Macroolithus) and E2 (comprising an unnamed ge­ nus). The present eggshell material differs from the ()otax~ andrewsi ZHAO, 1975, Mac­ roolithui; rugustus YOUNG, 1965 and Nanhsiungoo­ lith us chuetienensis ZHAO, 1975 in megascopic and microstructural characteristics. The overall shape in these three oospecies is obtuse at one end and pointed althe other with subcircular nodes and deep channeling (ZHAO, 1993); elongation ratios are also higher i.e. 2-2.2 in Elongatoolithus and 2-2.4 in Mac­ roolithus. The microstructure of Elongatoolithus an­ drewsi shows small mammillae with circular cruss sections and growth lines in upper spongy layer are almost straight. The other oospecies yaotunensis bears polar diameter 170 mm and shows tiny, parallel walled mammillae with indistinct margins. In comparison, in Ellipsoolithus a nearly oval shape is diagnostic and elongation ratio is 1.5-1.7 with polar diameter varying from 98-110 mm and Fig. 3 - Egg nest in plan and section; near Lavaria Mu­ equatorial diameterranging from 65-80 mm. The or­ wada locality, district Kheda, Gujarat; Lameta Formation; namentation comprises dispersituberculate to ra­ Upper Sandy Carbonate (calcretized paleosol); Late Cre­ motuberculate pattern at the polar end and taceous. lineartuberculate on the equatorial portion. Further the microstructure shows bulbous and nant components of Late Cretaceous assemblages fused mammillae with undulating grow1h line pattern of (ZHAO, 1975, 1994) and central Asian re­ in spongy layer; the spheroliths are characteristi­ gions of and China (KURZANOV & MIKHAI­ cally dome-shaped. LOV, 1989; MIKHAILOV, 1991, 1995; MIKHAILOV, SABATH & KURZANOV, 1994). PALAEOBIOLOGY In China, the oofamily Elongatoolithidae is stra­ Further, it is quite interesting to observe that both tigraphically associated with the Upper Cretaceous kind of eggshell materials (comprising those of ti­ Yuanpu Formation and Ping ling Formation exposed tanosaurids and the present eggshells) are re- in the Nanxiong Basin, though the reports from lower part of the Cretaceous exist (ZHAO, 1994). From these Upper Cretaceous formations, ZHAO (1975, 1994) described eggshells and nests of Macroo­ _ 10 lithus yaotunensis ZHAO, 1975, M. rugustus YOUNG, E 1965, Macroolithus sp., Elongatoolithus andrewsi ~ 9 ZHAO, 1975, E. elongatus YOUNG, 1954, Elongatoo­ .." lith us sp., and Nanhsiungoolithus chuetienensis ;; E o ZHAO, 1975. Of these, Elongatoolithus and Macroo­ .2 8 lith us were reassigned to elongatus, dis­ " covered by YOUNG (1965) based on material from .'§ 7 o Shantung Province, Guandung and Prov­ e 0 ~ 6 ince in China (MIKHAILOV, SABATH & KURZANOV, 0- W 1994 ). As far as the Central Asian material is concerned, 5 6 7 8 9 10 " 12 elongatoolithid eggshells are known from the Upper Polar diamet~r (em) Cretaceous Mongolian Barun Goyot, Djadokhta and Nemegtformations; only one locality is Lower Creta­ Fig. 4 - Plot of polar and equatorial diameter of ellipsoid ceous in age (MIKHAILOV, SABATH & KURZANOV, eggs, near Lavaria Muwada; district Kheda, Gujarat; 1994). This Mongolian eggshell material is catego­ Lameta Formation , Upper sandy Carbonate (calcretized rized into groups E1 (including Elongatoolithus and paleosol); Late Cretaceous.

383 R.S. LOYAL; D.M. MOHABEY; A. KHOSLA & A. SAHNI

Fig. 5 - Egg nests from near Lavaria Muwada; district Kheda, Gujarat; Lameta Formation, Upper Sandy Carbonate (calcretized paleosol); Upper Cretaceous. A - A nest showing seven ellipsoid shaped eggs (136IGSIIPAUCRlNGI94). B - A nest showing seven ellipsoid shaped eggs (137IGSIIPAUCRlNGI94). C - A nest showing three ellipsoid shaped eggs (138IGSIIPAU CRINGI94). D - Lineartuberculate ornamentation in equatorial region. E - Radial thin section show­ ing two layered eggshell microstructure showing dome-shaped spheroliths and tightly packed mammillary layer (149IGSIIPAUCRlNGI94). Bar (scale) represents A: 5cm; B: 5 cm; C: 5 cm; 0 : 1 cm; E: 250 ~m.

384 STATUS AND PALAEOBIOLOGY OF THE LATE CRETACEOUS INDIAN THEROPODS

External surface

Internal surface

Fig. 6 - Radial polished section of Eflipsoolithus khedaensis showing a single spherolith with a well developed mam­ millae (VPURl1 01); isolated eggshell at approximately 1.5 km NW of Rahioli, district Kheda, Gujarat; Lameta Formation; Upper Sandy Carbonate (calcretized paleosol); Upper Cretaceous. Bar (scale) represents 50 ~m.

385 R.S. LOYAL; D.M. MOHABEY; A. KHOSLA & A. SAHNI

stricted to the same lithology and also lie atthe same Delhi) for Young Scientist Scheme for 1998- stratigraphic level, suggesting important implica­ 2000 toAshu Khosla is gratefully acknowledged. tions on the community behaviour and dynamics of these i.e. a possible prey-predator relation­ ABBREVIATIONS ship. In this complex community scenario, though it CR -= 'Catalogue Register Number; GSI = Geo­ would be a bit premature to speculate on building a logical Survey of India; NG = Nagpur; PAL = Palae­ structured ichnoecoenoses (DODSON et a/., 1990) ontology Division + VPL = Vertebrate Palaeontology the present data points to abundant titanosaurid Laboratory; R=Rahioli populations providing a great predatory delight for the rarer abelisaurid and other theropods. REFERENCES BARSBOlD, R. & OSMOLSKA, H. (1990) - in CONCLUSIONS WEISHAMPEL, D.B.; DODSON , P. & OSM6LSKA, H. (Eds.), The The Late Cretaceous dinosaur fauna was domi­ DinosBuria, Univ. California Press, Berkeley, pp . 225-244,,- nated by the titanosaurids, abelisaurids and other BLANFORD, H.F. (1862) - On the Cretaceous and the other rocks of the South Areot and Tri chnopoly districts, South India. Mem. theropods. Though the megaloolithid eggs (broadly Geol. Surv.lndia, 4(1): 1-217. assignable to titanosaurids) occur in plenty along BONAPARTE, J.F. (1985) - A horned Cretaceous carnosaur from the east, west and central Narmada River region in Patagonia. Nat!. Geogr. Res., 1: 149-151 . the Lameta Formation and (MO­ BONAPARTE , J.F. & NOVAS , F.E. (1985)- Abelisaurus comahuen­ HABEY, 1983, 1984a, 1984b, 1986, 1990a, 1990b, sis, n.g" n.sp., del Cretaceo tardio de Patagonia. 1991, 1996a, 1996b; MOHABEY & MATHUR, 1989; Ameghiniana, 21 : 259-265. MOHABEY, UDHOJI & VERMA, 1993; KHOSLA, 1996; BUFFETAUT, E. (1987) - On the age of the dinosaur fauna from the Lameta Formation (Upper Cretaceous) of Central India. KHOSLA & SAHNI, 1995; SRIVASTAVA et a/., 1986; Newsl. Stratigr., 18: 1- 6. VIANEY-liAUD,JAIN & SAHNI, 1987; SAHNI, 1990, CHATTERJEE , S. (1978)-lndosuchusand Indosaurus, Cretaceous 1993; SAHNI et a/., 1994) and seven to eight of its 00- carnosaurs from India. J. Palaeontol., 52(3) : 570-580. species are known (SAHNI & KHOSLA, 1994; KHO­ CHATTERJEE, S. & CREISLER, B.S. (1994) - Alwalkeria (Theropo­ SLA, 1996; KHOSLA & SAHNI, 1995; MOHABEY, da) and Mortuneria (Plesiosauria), new names for preoccu­ 1996a; LOYAL, KHOSLA & SAHNI, 1996), the elonga­ pied Walkeria CHATTERJEE, 1987 and Turneria CHATERJEE & toolithid eggs (assignable to theropods) are SMALL , 1989. J. Vertebr. Paleontol., 14(1): 142. scarcely known excepting the present find of Ellip­ CHATTERJEE, S. & RUDRA, O.K. (1996) - KT events in India: im­ pact, rifting, volca nism and dinosaur extinction. Mem. Que­ soolithus (MOHABEY, 1996b) from a nesting site at ensl. Museum, 39(3): 489-532. Lawaria Muwada and Kevadiya (near Rahioli). Fur­ DODSON , P.; COOMBS , W.P.; FARLOW, J.D. & TATARINOV, L.P. ther, the present collection comprises only (1990) - Dinosaur Paleobiology in WEISHAMPEl, D.B.; DOD­ eggshells, nests, isolated eggs and shell debris; un­ SON, P. & DSMOLSKA, H. (Eds.), The Dinosauria, Univ. Califor­ fortunately no associated skeletallembryonic mate­ nia Press, Berkeley, pp. 31-62. rial was found, which could warrant proper HUENE, FV. & MATLEY, C.A. (1933) - The Cretaceous and Ornithischia of the central provinces of India. Mem. Geol. affiliations of these eggshells to the generic level. In Surv. India. Palaeontol. Indica, 21(1 ): 1-72. this context, we are of the opinion that the abeli­ KHOSLA, A. & SAHNI , A. (1995) - Parataxonomic classification of saurids could have been a dominant Indian Late Late Cretaceous dinosaur eggshells from India . J. Palaeontol. Cretaceous dinosaur group; similar view on abeli­ Soc. India, 40: 87-102. saurid distribution is expressed by CHATTERJEE & KHOSLA, A. (1996) - Dinosaur eggshells from the Late Cretaceous RUDRA (1996). We, henceforth, urgently require Lameta Formation along the east-central Narbada River Re­ gion : Biomineralization and Morphotaxonomical Studies. more field discoveries of elongatoolithid eggshells Ph.D. Thesis, Panjab Univ., 197 pp. (unpublished). with associated skeletal material as also the track­ KRAUSE, D.W. & DODSON, P. (1994) - The premaxilla of Majunga­ way evidences, not only from the Indian landmass saurus (Theropoda), Late Cretaceous, Madagascar: implica­ but also from biogeographically contiguous Mada­ tions for relationships. J. Verlebr. Paleonto/., 14: 32A. gascar region. KRAUSE, D.W. & HARTMAN, J.H. (1996) - Late Cretaceous foss ils from Madagascar and their implications for Biogeographic Relationships with the Indian Subcontinent, in SAHNI , A. (Ed.), ACKNOWLEDGEMENTS Cretaceous Stratigraphy and Palaeoenvironments - Rama Rao Volume. Mem. Geol. Soc. India, 37: 135-154. We are thankful to Profs. Zhao Zikui (China) and KURZANOV, S.M. & MIKHAILOV, K.E. (1989) - Dinosaur eggshells Konstantin E. Mikhailov (Russia) for sending litera­ from the Lower Cretaceous of Mongolia, in GILLETTE , D.O. & ture and providing information on Cretaceous egg­ LOCKLEY, M.G. (Eds.), Dinosaur Tra cks and Traces , Cam­ shell material. Financial assistance in the form of bridge Univ. Press, New York, pp. 109-113. Senior Research Fellow (No. 9/135/(287)/95/EM R­ LOYAL, RS.: KHO SLA, A. & SAHNI , A. (1996) - Gondwanan dino­ I) awarded by the Council of Scientific and Industrial saurs of India: affinities and palaeobiogeography. Mem. Que­ ensl. Museum, 39(3): 627-328. Research (CSIR) New Delhi; CIES (Paris, France) LYDEKKER , R (1879) - Indian Pre-Tertiary Vertebrata. Fossil Rep­ for postdoctoral fellowship during years 1997-1998 tilia and Batrachia. Palaeontol.lndica , Ser. 4, 1(3): 1-36. and Department of Science and Technology (New

386 STA TUS AND PALAEOBIOLOGY OF THE LATE CRETACEOUS INDIAN THEROPODS

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