GAIA N' 15, LlSBOAlLISBON, DEZEMBRO/DECEMBER 1998, pp. 379-387 (ISSN: 0871-5424) STATUS AND PALAEOBIOLOGY OF THE LATE CRETACEOUS INDIAN THEROPODS WITH DESCRIPTION OF A NEW THEROPOD EGGSHELL OOGENUS AND OOSPECIES, ELLIPSOOLITHUS KHEDAENSIS, FROM THE LAMETA FORMATION, DISTRICT KHEDA, GUJARAT, WESTERN INDIA 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 Abelisauridae; recent reports confirm the presence of Majungasaurus crenatissimus (DEPRET, 1896), Indosuchus HUENE & MATLEY, 1933 possible Carnotaurus BONAPARTE, 1985 and ?Allosaurus 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, Laevisuchus 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 dinosaur eggshell material referable to a new ooge­ nus and oospecies Ellipsoolithus khedaensis, (oofamily Elongatoolithidae) from near Rahioli, district Kheda, Gujarat. This material has been presently assigned to the Theropoda and can be easily differentiated from the other well known megaloolithid oospecies (broadly referable to Sauropoda 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 taxonomy 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 clade Abelisauridae. 1933; CHATTERJEE, 1978; CHATTERJEE & RUDRA, This viewpoint is also corroborated by CHATTERJEE 1996). Except for the presence of a Triassic 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 Jurassic 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 Allosauridae (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 LATE CRETACEOUS 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