India's First Dinosaur, Rediscovered

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Geological Society of India, Bangalore, physics of the Precambrians (ed. Saha, 14. Mondal, N., Problems of superposed Memoir 27, 1994. A. K.), Hindustan Publications, New folding: An experimental study, unpub- 3. Sarkar, S. N. and Saha, A. K., Q. J. Delhi, 1996, pp. 1–14. lished Ph D thesis, Jadavpur University, Geol., Min. Metall. Soc. India, 1962, 34, 10. Mazumdar, S. K., Indian Miner., 1996, 1991. 97–136. 93, 139–174. 4. Sarkar, S. N. and Saha, A. K., Geol. 11. Gupta, A. and Basu, A., North Singhb- Received 12 September 2011; re-revised Mag., 1963, 100, 69–92. hum Proterozoic Mobile Belt, Eastern accepted 20 November 2012 5. Naha, K., Sci. Cult., 1956, 22, 43–45. India – A Review, M. S. Krishnan Cen- 6. Naha, K., Geol. Mag., 1959, 96, 137–140. tenary Volume, Geol. Surv. India Spec. ALOKESH CHATTERJEE* 7. Naha, K., Q. J. Geol., Min. Metall. Soc. Publ. No. 55, 2000, pp. 195–226. SHARMILA BHATTACHARYA India, 1965, 37, 41–95. 12. Haneberg, W. C., Cuspate–lobate folds

8. Bhattacharya, D. S. and Sanyal, P., In along a sedimentary contact, Los Lunas Department of Geology, Precambrian of the Eastern Indian Volcano, New Mexico. New Mexico Shield (ed. Mukhopadhyay, D.), Geo- Bureau of Mines and Mineral Resources Presidency University, logical Society of India, Bangalore, Bulletin 137, 1991, pp. 162–163. 86/1 College Street, Memoir 8, 1988, pp. 85–111. 13. Ghosh, S. K., Structural Geology Fun- Kolkata 700 073, India 9. Pradhan, A. K. and Srivastava, D. C., In damentals and Modern Developments, *For correspondence. Recent Researches in Geology and Geo- Pergamon Press, 1993. e-mail: [email protected]

India’s first dinosaur, rediscovered

‘Reasoning from analogy at Jubbulpore, sented illustrations of the better preserved whose diagnostic features were eventu- where some of the basaltic cappings of element4. Falconer correctly identified ally made obsolete by the discovery of the hills had evidently been thrown out of the vertebrae as reptilian but refrained more complete remains12,13. The first craters long after this surface had been from coining a name for them, probably such discoveries were made by Charles raised above the waters, and become the because the manuscript, published post- Matley and Durgasankar Bhattacharji in habitation both of vegetable and animal humously, was not intended for publica- the early 1900s (refs 14 and 15). Excava- life, I made the first discovery of fossil tion. It was not until 1877 that Sleeman’s tions near the probable location of Slee- remains in the Nerbudda valley. I went discovery was finally recognized as a man’s original site in Jabalpur produced first to a hill within sight of my house in new genus and species of sauropod dino- the braincase and partial skeleton of Ant- 1828, and searched exactly between the saur called Titanosaurus indicus (‘Indian arctosaurus septentrionalis16 (whose plateau of basalt that covered it, and the titan reptile’) by Richard Lydekker5. He genus name has since been changed to stratum immediately below; and there I named a second species from Jabalpur, Jainosaurus17), a partial postcranial found several small trees with roots, T. blanfordi (‘Blanford’s titan reptile’), skeleton of a second, smaller individual trunks, and branches, all entire, and just two years later6. At that time, only of the same genus18,19, as well as many beautifully petrified. They had been only approximately 115 dinosaur species had isolated bones. Many of the theropod recently uncovered by the washing away been identified, less than 10% of the remains were shipped to London for of a part of the basaltic plateau. I soon 1401 species known by 2004 (ref. 7). preparation and description in 1922 and after found some fossil bones of ani- After passing safely through many 1925; most of them were returned to mals.’ hands for over the course of half a cen- India in 1936, along with a plaster cast of –W. H. Sleeman1 tury, the original remains of T. indicus the partial hind limb of Jainosaurus2. went missing. It is not known exactly There is no manifest for this shipment, so So begins the history of Indian dinosaur when this happened, or even who was the it is not known exactly which specimens studies. The bones that Sleeman col- last to examine them, but we were not made the trip to and from London. Bar- lected from the beds underlying the Dec- able to find mention of first-hand obser- num Brown visited Bara Simla and made can Traps at Jabalpur would soon embark vation of T. indicus bones subsequent to additional collections of theropod20 and on ‘rambles’ of their own2. Sleeman sent Lydekker’s last treatment of them6. A sauropod21,22 remains, which are housed two fossil bones to G. G. Spilsbury, a cast of the specimen is present in the at the American Museum of Natural His- civil surgeon, who himself collected a Natural History Museum, London tory. More recent excavations elsewhere third bone from the same bed. In 1832, (NHMUK OR40867). in India have added to the initial discov- Spilsbury sent all three to Calcutta anti- T. indicus holds a special place as eries of Matley and Bhattacharji at Bara quarian, James Prinsep. By 1862, these India’s first recorded dinosaur, discov- Simla and Chhota Simla (Jabalpur). One fossil bones were presented to Thomas ered only four years after the discovery of most important was an accumulation Oldham, the first Director of the Geo- of the first-named dinosaur Megalosau- of several hundred bones21,23–25 in strata logical Survey of India. Surgeon– rus8 and 14 years before the name ‘Dino- just below a prolific egg-bearing hori- botanist Hugh Falconer, co-discoverer of sauria’ was coined9. Like Megalosaurus zon26,27 in Rahioli, western India. An- the Siwalik vertebrates3, provided the and other early-named dinosaurs such as other such locality was Dongargaon in first description of two bones, which he Iguanodon10 and Cetiosaurus11, Titano- central India, some 335 km south of identified as caudal vertebrae, and pre- saurus was based on limited material Jabalpur. The Dongargaon locality

34 CURRENT SCIENCE, VOL. 104, NO. 1, 10 JANUARY 2013 SCIENTIFIC CORRESPONDENCE produced the most complete skeleton of skulls bearing narrow tooth crowns36,37, nity to access and study T. indicus is an Indian dinosaur, Isisaurus colberti presacral vertebrae with complex lamina- symptomatic of a larger issue. There are (formerly known as ‘Titanosaurus’ col- tion38 and limbs that were slightly angled several Indian dinosaur specimens that berti), which is known from a braincase, outward in a wide-gauge posture39. Some are currently missing, including both presacral, sacral and caudal vertebrae, titanosaurs even possessed dermal small and large specimens of sauropod girdle bones and limb bones28,29. The armour22,40 that may have functioned as a and theropod dinosaurs. Notable missing more recent discovery of scores of dino- mineral store that allowed them to sur- specimens include the partial postcranial saur bones in Balochistan, Pakistan30–32 vive in stressed environments41. In addi- skeleton of the stocky-limbed, large also represents an important source of tion, Titanosaurus provided the first theropod Lametasaurus indicus44, skull information on dinosaurs of the Indian indications of what Lydekker42 called a materials of both Indosaurus matleyi and subcontinent. ‘remarkable community of type which Indosuchus raptorius, parts of Jainosau- Beyond its historical and patrimonial undoubtedly exists between the faunas of rus septentrionalis and the small no- significance, what is the relevance of T. southern continents of the world’. Recast asaurid theropod Laevisuchus indicus indicus, if the species was based on lim- in today’s mobilist palaeogeographic and many theropod limb bones16 (Figure ited material, now missing, deemed paradigm, the Indian subcontinent takes 1). The nonavailability of these elements insufficient to distinguish it from other on special significance as a large disper- has seriously hindered efforts to under- dinosaurs13? The material is relevant for sal vector43 that began the Mesozoic stand the evolutionary history of Indian several reasons. First, Titanosaurus pro- interlocked with other southern land- dinosaurs and to decode their palaeobio- vided an initial glimpse at, and eventually masses and ended it in isolation prior to geographic connections to other southern became the namesake for, the diverse, docking on Asia sometime in the early landmasses45,46. But are these bones lost, late-surviving sauropod lineage Titano- Tertiary. Large continental tetrapods like or merely misplaced? Are efforts best sauria33. Titanosaurs comprise more than Titanosaurus and other dinosaurs can directed at retrieving these bones in col- 40 genera34, which have been recorded provide insight into India’s relationship lections or finding new bones in the from all continental landmasses, includ- with other landmasses, both southern and field? ing Antarctica35. They are morphologi- northern. Last, and perhaps most signifi- The Geological Survey of India (GSI) cally distinctive sauropods with elongate cantly, inability of the scientific commu- and the University of Michigan have recently embarked on a programme to recover missing fossil bones in museum collections and to collect new bones from field sites. Efforts at the Indian Museum (Kolkata) and GSI repositories have resulted in the recovery of the misplaced holotypic caudal vertebra of T. indicus (Figure 2). The T. indicus holotype was stored together with bones of T. blanfordi and with Triassic vertebrates of the ‘Lydekker collection’47, also pre- sumed missing. The bones were recov- ered from the vast fossil vertebrate and

Figure 2. Rediscovered holotypic caudal Figure 1. Currently missing Indian dinosaur fossils. a, b, Syntypic skull elements of the large vertebra of India’s first dinosaur, Titanosau- theropod Indosuchus raptorius. a, Skull roof K20/350 in dorsal view. b, Skull roof and braincase rus indicus, in left lateral view. Roman K27/685 in dorsal and left lateral views16. c, Holotypic partial skeleton of the large theropod numerals inked on the bone refer to plate and dinosaur Lametasaurus indicus, including sacrum in ventrolateral view, right ilium in ventral figure numbers; the other two numbers (GSI view (anterior towards bottom), and left tibia in lateral view44. d, Holotypic cervical vertebra ‘2191’, ‘2194’) represent serial numbers as (K20/614) of the noasaurid Laevisuchus indicus16. Scale equals 10 cm for (a) and (b), 30 cm for recorded in the Specimen Register of the (c) and 5 cm for (d). Curatorial Division, GSI. Scale equals 5 cm.

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3 – no accession number, stored outside the ‘normal’ area – suggest that future rediscoveries may rely on visual recognition of unlabelled specimens that may be stored apart from other collections. These results emphasize the impor- tance of fossil repositories as secure storage for historical objects that form the basis for scientific research. These objects constitute the primary record of the evolutionary history of Greater India and its past and present connections to other landmasses.

1. Sleeman, W. H., Rambles and Recollec- tions of an Indian Official, J. Hatchard and Son, London, 1844, vol. 1, p. 478. 2. Carrano, M. T., Wilson, J. A. and Bar- rett, P. M., In Dinosaurs and other Ex- tinct Saurians: A Historical Perspective (eds Moody, R. T. J. et al.), Geological Society, London, Special Publication, 2010, pp. 161–173. 3. Nair, S. P., Sci. Context, 2005, 18, 359–

392. Figure 3. Other rediscovered Indian dinosaur specimens. a, Abelisaurid femur (GSI K27/569) in anterior view. b, Holotypic cervical vertebra of Laevisuchus indicus (GSI K20/613) in left lat- 4. Falconer, H., In Palaeontological Mem- eral view. c, Cast of Jainosaurus cf. septentrionalis hind limb from Chhota Simla18,19, presented oirs and Notes of the late Hugh Fal- by Natural History Museum (London) to Nagpur City Museum2 in 1936. The original specimens coner, Vol. 1, Fauna Antiqua Sivalensis are labelled as NHMUK 5903. d, Undescribed noasaurid caudal vertebra from the Matley Collec- (ed. Murchison, C.), Robert Hardwicke, tion (GSI K20/612) in right lateral view. e, Holotypic caudal vertebra of Titanosaurus blanfordi London, 1868, pp. 414–416 (written in (GSI 2195) in right lateral view. Scale bar equals 10 cm for (a), 2 cm for (b), 30 cm for (c), 1862 and published posthumously). 1.5 cm for (d) and 5 cm for (e). 5. Lydekker, R., Rec. Geol. Surv. India, 1877, 10, 30–43. invertebrate collection of the Curatorial a collection of theropod cranial, caudal, 6. Lydekker, R., Palaeontol. Indica (Ser. 4), 1879, 1, 20–33. Division of GSI Headquarters at Kolkata. and limb elements collected by Matley 7. Benton, M. J., Biol. Lett., 2008, 6, 544– The T. indicus caudal vertebra is pre- but never described was found, along 547. served intact, based on comparisons with with undescribed rib fragments of the 8. Buckland, W., Trans. Geol. Soc. London, 4,5 19 drawings by Falconer and Lydekker , Chhota Simla specimen of Jainosaurus , 1824, 21, 390–397. save a small portion of its cotylar rim still in their original wrappings. In each 9. Owen, R., Rep. Br. Assoc. Adv. Sci., that is now broken. Recovery efforts of these instances, the specimen had no 1842, 1841, 60–204. have also turned up several other bones accession number – either because it never 10. Mantell, G., Philos. Trans. R. Soc. Lon- (Figure 3). The original syntypic caudal received one (e.g. T. indicus, T. blan- don, 1825, 115, 179–186. vertebrae of T. blanfordi (one of which fordi) or because the number had been 11. Owen, R., Proc. Geol. Soc. London, later was removed from the type16) were separated from it by breakage (e.g. 1841, 3, 457–462. 12. Sues, H.-D., Bull. Zool. Nomencl., 1998, found with the T. indicus holotype. The Laevisuchus, Jainosaurus). Recovery of 55, 240–241. holotypic humerus of J. septentrionalis these specimens was more difficult 13. Wilson, J. A. and Upchurch, P., J. Syst. (GSI K27/497) was found in several because individual specimens (often Palaeontol., 2003, 1, 125–160. pieces atop tall display cases in the Si- fragments), rather than numbers, had to 14. Matley, C. A., Rec. Geol. Soc. India, walik Gallery of the Indian Museum after be recognized. In several cases (e.g. T. 1921, 53, 142–164. having gone unnoticed for decades48. indicus, T. blanfordi, Jainosaurus), the 15. Mohabey, D. M., J. Palaeontol. Soc. One of the holotypic cervical vertebrae specimen had been shifted outside the India, 2011, 56, 127–135. of the small theropod Laevisuchus indi- designated collection space, which had 16. Huene, F. V. and Matley, C. A., Palae- cus (GSI K20/613)16 was recently found been searched repeatedly. ontol. Indica, 1933, 21, 1–74. in three separate pieces in unmarked Recovery of T. indicus and other mis- 17. Hunt, A. P., Lockley, M. G., Lucas, S. G. and Meyer, C. A., Gaia, 1994, 10, 261– boxes, together with other unnumbered placed fossils bodes well for retrieval of 279. fragments in the Invertebrate Gallery of other important missing specimens, such 18. Swinton, W. E., Ann. Mag. Nat. Hist. the Indian Museum. A complete abeli- as Lametasaurus indicus, Indosaurus (Ser. 11), 1947, 14, 112–123. 16 saurid femur (GSI K27/569) was found matleyi and Indosuchus raptorius (Figure 19. Wilson, J. A., Barrett, P. M. and Car- in five pieces in the Siwalik Gallery of 1). The circumstances associated with rano, M. T., Palaeontology, 2011, 54, the Indian Museum. In that same cabinet, the loss of the bones in Figures 2 and 981–998.

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20. Chatterjee, S., J. Palaeontol., 1978, 52, 36. Nowinski, A., Palaeontol. Pol., 1971, Museum of Paleontology, under a Memoran- 570–580. 25, 57–81. dum of Understanding for study of Late Cre- 21. Chatterjee, S. and Rudra, D. K., Mem. 37. Zaher, H. et al., PLoS ONE, 2011, 6, taceous tetrapod fossils from the Lameta Queensl. Mus., 1996, 39, 489–532. e16663. Formation. For technical and logistic support, 22. D’Emic, M. D., Wilson, J. A. and Chat- 38. Salgado, L. and Powell, J. E., J. Vertebr. we thank A. Sundaramoorty, Director Gen- terjee, S., J. Vertebr. Palaeontol., 2009, Palaeontol., 2010, 30, 1760–1772. eral, GSI; Dr K. Rajaram, Deputy Director 29, 165–177. 39. Wilson, J. A. and Carrano, M. T., Paleo- General, PSS (retired); P. Sarkar, Director, 23. Mathur, U. B. and Pant, S. C., J. Palae- biology, 1999, 25, 252–267. Curatorial Division, and D. V. R. Ramna ontol. Soc. India, 1986, 31, 22–25. 40. Bonaparte, J. F. and Powell, J. E., Mem. Murthy, Deputy Director General, GSI, 24. Mathur, U. B. and Srivastava, S., J. Geol. Soc. Géol. Fr. (Nouvelle Sér.), 1980, Lucknow. J.A.W. was supported by grants Soc. India, 1987, 29, 554–566. 139, 19–28. from the National Geographic Society Com- 25. Dwivedi, G. N., Mohabey, D. M. and 41. Curry Rogers, K. A., D’Emic, M. D., mittee for Research and Exploration Grant Bandyopadhyay, S., Curr. Trends Geol., Rogers, R. R., Vickaryous, M. and Cagan, 8127-06, the Jurassic Foundation, and a Sen- 1982, 7, 79–87. A., Nature Commun., 2011, 2, 564–569. ior Fellowship from the American Institute of 26. Mohabey, D. M., J. Geol. Soc. India, 42. Lydekker, R., An. Mus. Plata, 1893, 2, Indian Studies. 1984, 25, 329–337. 1–14. 27. Srivastava, S., Mohabey, D. M., Sahni, 43. McKenna, M. C., In Implications of Con- Received 3 October 2012; accepted 15 A. and Pant, S. C., Palaeontogr., Abt. A, tinental Drift to the Earth Sciences (eds November 2012 1986, 193, 219–233. Tarling, D. H. and Runcorn, S. K.), Aca- 28. Berman, D. S. and Jain, S. L., Ann. Car- demic Press, London, 1973, pp. 295–308. 1 negie Mus., 1982, 51, 405–422. 44. Matley, C. A., Rec. Geol. Surv. India, DHANANJAY M. MOHABEY 2 29. Jain, S. L. and Bandyopadhyay, S., J. 1924, 55, 105–109. SUBASHIS SEN 3, Vertebr. Palaeontol., 1997, 17, 114–136. 45. Novas, F. E., Agnolin, F. L. and Bandyo- JEFFREY A. WILSON * 30. Malkani, M. S. and Anwar, C. M., Geol. padhyay, S., Rev. Mus. Argent. Cienc. Surv. Pak. Inf. Release, 2000, 732, 1–16. Nat., Nuevo Ser., 2004, 6, 67–103. 1 31. Wilson, J. A., Malkani, M. S. and Gin- 46. Carrano, M. T., Sertich, J. J. W. and Geological Survey of India gerich, P. D., Gondwana Geol. Mag., Loewen, M. A., Smithsonian Contrib. (Northern Region), 2005, 8, 101–109. Paleobiol., 2011, 35, 1–53. Lucknow 226 004, India 32. Malkani, M. S., J. Appl. Emerg. Sci., 47. Lydekker, R., Palaeontol. Indica (Ser. 2Geological Survey of India 2006, 1, 108–140. 4), 1885, 1, 1–38. (Central Headquarters), 33. Bonaparte, J. F. and Coria, R. A., 48. Wilson, J. A., D’Emic, M. D., Curry Kolkata 700 016, India Ameghiniana, 1993, 30, 271–282. Rogers, K. A., Mohabey, D. M. and Sen, 3Museum of Paleontology and 34. Wilson, J. A., Actas de las III Jornadas S., Contrib. Mus. Paleontol. Univ. Mich., Department of Earth and Internacionales sobre Paleontología de 2009, 32, 17–40. Environmental Sciences, Dinosaurios y su entorno. Salas de los Infantes, Burgos, 2006, pp. 169–190. University of Michigan, 35. Cerda, I. A., Carabajal, A. P., Salgado, ACKNOWLEDGEMENTS. The finding pre- Ann Arbor, L., Coria, R. A., Reguero, M. A., Tam- sented is the result of a joint collaborative Michigan 48109-1079, USA bussi, C. P. and Moly, J. J., Naturwissen- programme involving the Geological Survey *For correspondence. schaften, 2011, 99, 83–87. of India (GSI) and University of Michigan e-mail: [email protected]

Mass stranding of pilot whale Globicephala macrorhynchus Gray, 1846 in North Andaman coast

Pilot whale is a carnivorous marine and long-finned whales is similar. How- and Pacific Oceans. Both the species live mammal described under the order Ceta- ever, the fin of the long-finned whales is in groups of 20–60 individuals or more. cea, suborder Odontoceti (toothed whales). one-fifth or more of their body length The population of G. macrorhynchus has Though commonly called as ‘black fish’ and one-sixth for that of short-finned been estimated as 150,000 in the eastern or ‘pothead whales’, these mammals are whales. Short-finned pilot whales have tropical Pacific Ocean and about 30,000 named as ‘pilot whales’ because it was fewer teeth, i.e. 7–9 short, sharply in the western Pacific, off the coast of believed that pods were piloted by a pointed teeth in the front of each tooth Japan2. Normally they prefer the waters leader1,2. They are gregarious and fre- row, whereas it is 8–13 for long-finned of the shelf break and slope2. Although quently found with other small cetace- whales3. According to IUCN Red List, they primarily feed on squid4, pilot ans. Pilot whales are one of the largest both the species are insufficiently whales consume fishes, including Atlan- members of the family Delphinidae. Two known. Pilot whales are found in waters tic cod, Greenland turbot, Atlantic mack- extant species of pilot whales reported in nearly worldwide with long-finned pilot erel, Atlantic herring, hake, blue whiting the world oceans are long-finned Globi- whales living in temperate waters, and and spiny dogfish2,5. These whales are cephala melas (Traill, 1809) and short- short-finned pilot whales living in the habituated to migrate seasonally inshore finned Globicephala macrorhynchus tropical and subtropical waters generally and offshore in response to the dispersal Gray 1846. General appearance of short- in deep offshore areas of Indian, Atlantic of their prey2. Pilot whales are often

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