• ARCHANA SHARMA Geological Survey of India image/svg+xml
  • FERNANDO E. NOVAS Laboratorio de Anatomía Comparada y Evolución de los Vertebrados CONICET–Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Buenos Aires, Argentina
  • SANJAY SINGH Geological Survey of India image/svg+xml



Pedal ungual phalanx; Spinosauridae; Jaisalmer Basin; Middle Jurassic; Bathonian; Eastern Gondwana.


We describe an isolated, almost complete pedal ungual phalanx from the Middle Jurassic marine carbonate rocks of the Jaisalmer Basin, Rajasthan, north-western India. The ungual bone is triangular shaped, pointed, elongated, asymmetrical, dorsoventrally compressed, ventrally flat, bearing a shallow semi-circular excavation, and almost straight in lateral view. The morphological features, as well as its proportions closely resemble the pedal ungual phalanges of spinosaurid theropods, presently known dominantly from the Cretaceous. The affinity to spinosaurids is supported by bivariate and multivariate analyses. The ungual phalanx is tentatively identified as a basally branching Jurassic spinosaurid under Megalosauroidea. Considering the stratigraphical and geographical provenances, this contribution may represent the oldest record of a spinosaurid.


Metrics Loading ...

Author Biography

ARCHANA SHARMA, Geological Survey of India

Post: Director

Western Region

State Unit Rajasthan

Geological Survey of India, Jaipur


Agassiz L.J.R. (1838) - Recherches sur les poissons fossiles, Vol. 3. Imprimerie de Petitpierre, Neuchâtel.

Allain R. & Chure D. (2002) - Poekilopleuron bucklandii, the theropod dinosaur from the Middle Jurassic (Bathonian) of Normandy. Palaeontology, 45(6): 1107-1121.

Allain R., Xaisanavong T., Richir P. & Khentavong B. (2012) - The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the Early Cretaceous of Laos. Naturwissenschaften, 99: 369-377.

Ahmad F., Quasim M.A., Ghaznavi M.A., Khan Z. & Ahmad A.H.M. (2017) - Depositional Environment of the Fort Member of the Jurassic Jaisalmer Formation, (Western Rajasthan, India) as revealed from Litho Facies and Grain Size Analysis. Geological Acta, 15(3): 153-167.

Averianov A.O., Krasnolutskii S.A. & Ivantsov S.V. (2010) - A new basal coelurosaur (Dinosauria: Theropoda) from the Middle Jurassic of Siberia. Proceedings of the Zoological Institute, 314(1): 42-57.

Barker C.T., Hone D.W.E., Naish D., Cau A., Lockwood J.A.F., Foster B., Clarkin C.E., Schneider P. & Gostling N.J. (2021) - New spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae. Scientific Reports, 11(1):19340.

Becerra M., Pol D., Rauhut O. & Cerda I. (2016) - New Heterodontosaurid remains from the Cañadón Asfalto Formation: cursoriality and the functional importance of the pes in small Heterodontosaurids. Journal of Paleontology, 1: 1-23.

Benson R.B.J. (2010) - A description of Megalosaurus bucklandii (Dinosauria, Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society, 158: 882-935.

Bertozzo F., Dalla Vecchia F.M. & Fabbri M. (2017) - The Venice specimen of Ouranosaurus nigeriensis (Dinosauria, Ornithopoda). PeerJ, 5: e3403.

Blanford W.T. (1877) - Geological notes on the Great Indian Desert between Sind and Rajputana. Record Geological Survey of India, 10(Pt. 1): 10-21.

Brochu C.A. (2003) - Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology. 22: 1-138.

Brown C.M., Boyd C.A. & Russell A.P. (2011) - A new basal ornithopod dinosaur (Frenchman Formation, Saskatchewan, Canada), and implications for late Maastrichtian ornithischian diversity in North America. Zoological Journal of the Linnean Society, 163(4): 1157-1198.

Brownstein C.D. (2017) - Description of Arundel Clay ornithomimosaur material and a reinterpretation of Nedcolbertia justinhofmanni as an “Ostrich Dinosaur”: Biogeographic Implications. Peerj, 5: E3110, 1-20.

Brusatte S., Benson B.J. & Xing X. (2010) - The evolution of large-bodied theropod dinosaurs during the Mesozoic in Asia. Journal of Iberian Geology. 36(2): 275-296.

Buffetaut E. (2008) - Spinosaurid teeth from the Late Jurassic of Tengaduru, Tanzania, with remarks on the evolutionary and biogeographical history of the Spinosauridae. In: Mazin J.M., Pouch J., Hantzpergue P. & Lacombe V. (Eds.) - Mid–Mesozoic Life and Environments. Lyon: Documents Des Laboratoires De Géologie Lyon: 26-28.

Carpenter K. & Galton P.M. (2018) - A photo documentation of bipedal ornithischian dinosaurs from the Upper Jurassic Morrison Formation, USA. Geology of the Intermountain West, 5: 167-207.

Carrano M.T. (2007) - The appendicular skeleton of Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. In: Sampson S.D. & Krause D.W. (Eds.) - Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology. Society of Vertebrate Paleontology Memoir, 8(27): 163-179.

Carrano M.T., Benson R.B. & Sampson S.D. (2012) - The Phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology, 10: 211-300.

Carrano M.T., Sampson S.D. & Forster C.A. (2002) - The osteology of Masiakasaurus knopfleri, a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology, 22: 510-534.

Carter H.J. (1861) - On the Contribution to the Geology of Western India Including Sind and Baluchistan. Journal Royal Asiatic Society, 6(21): 161-206.

Chanthasit P. (2010) - The ornithopod dinosaur Rhabdodon from the Late Cretaceous of France: anatomy, systematics and paleobiology. Université Claude Bernard, Lyon I. English. NNT: 2010LYO10098.

Charig A.J. & Milner A.C. (1986) - Baryonyx, a remarkable new theropod dinosaur. Nature, 324(6095): 359-361.

Charig A.J. & Milner A.C. (1997) - Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey. Bulletin of the Natural History Museum of London, 53: 11-70.

Colbert E.H. (1962). Dinosaurs: their discovery and their World. 288 pp., London.

Cruzado-Caballero P., Díaz-Martínez I., Rothschild B., Bedell M. & Pereda-Suberbiola X. (2021) - A limping dinosaur in the Late Jurassic: Pathologies in the pes of the neornithischian Othnielosaurus consors from the Morrison Formation (Upper Jurassic, USA), Historical Biology, 33 (9): 1753-1759.

Currie P.J. & Zhao X. (1993) - A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjiang, People’s Republic of China. Canadian Journal of Earth Sciences, 30(10): 2037-2081.

Das Gupta S.K. (1975) - A Revision of the Mesozoic-Tertiary Stratigraphy of the Jaisalmer Basin, Rajasthan. Indian Journal of Earth Sciences 2(1): 77-94.

De França T., Santos B.N., De Oliveira M.N.Y, Alfredo M.M., Matos L.R. & Rodrigues C.F. (2021) - The first record of a spinosaurid pedal ungual from Brazil (Boca Do Forno Ravine, Itapecuru Formation, Parnaı´Ba Basin). Historical Biology, 34(9): 1817-1826.

De Lapparent A.F. (1960) - Les dinosauriens du “continental intercalaire” du Sahara central. Mémoires de la Société géologique de France, 88(A): 1-57.

Dieudonné P., Tortosa T., Torcida Fernández-Baldor F., Canudo J.I. & Díaz-Martínez I. (2016) - An unexpected early rhabdodontid from Europe (Lower Cretaceous of Salas de los Infantes, Burgos Province, Spain) and a re-examination of basal iguanodontian relationships. PLoS ONE, 11(6): e0156251.

Fitzinger L. (1843) - Systema Reptilium. Fasciculus Primus. Amblyglossae. Apud Braumüller and Seidel Bibliopolas, Vienna.

Forster C.A. (1990) - The postcranial skeleton of the ornithopod dinosaur Tenontosaurus tilletti. Journal of Vertebrate Paleontology, 10(3): 273-294.

Galton P.M. (1981) - Dryosaurus, a hypsilophodontid dinosaur from the Upper Jurassic of North America and Africa Postcranial skeleton. Paläontologische Zeitschrift, 55: 271-312.

Galton P. (2009) - Notes on Neocomian (Lower Cretaceous) Ornithopod dinosaurs from England - Hypsilophodon, Valdosaurus, “Camptosaurus”, “Iguanodon” - and referred specimens from Romania and elsewhere. Revue De Paleobiologie, 28: 211-273.

Galton P. & Powell H.P. (1980) - The ornithischian dinosaur Camptosaurus prestwichii from the Upper Jurassic of England. Palaeontology, 23: 411-443.

Gauthier J. (1986) - Saurischian monophyly and the origin of birds. Memoirs of the California Academy of Sciences, 8: 1-55.

Gilmore C.W. (1909). Osteology of the Jurassic reptile Camptosaurus, with a revision of the species of the genus, and descriptions of two new species. Proceedings of the United States National Museum, 36: 197-332.

Gilmore C.W. (1915) - Osteology of Thescelosaurus, an ornithopodous dinosaur from the Lance Formation of Wyoming. Proceedings of the United States National Museum, 49(2127): 591-616.

Hammer Ø., Harper D.A.T. & Paul D.R. (2001) - Past: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontologia Electronica, 4(1), 4: 9 pp.

He X. & Cai K. (1984) - The Middle Jurassic dinosaurian fauna from Dashanpu, Zigong, Sichuan Vol. I - The ornithopod dinosaurs. Sichuan Scientific and Technological Publishing House Chengdu, 71 pp. [In Chinese with English Summary].

Hitchcock E. (1836) - Ornithichnology. Description of the footmarks of birds, (Ornithichnites) on New Red Sandstone in Massachusetts. American Journal of Science, 29(ser. 1): 307-340.

Hitchcock E. (1858) - Ichnology 01 New England. A report on the sandstone of the Connecticut valley, especially its fossil footmarks. William White, Boston.

Hocknull S.A., White M.A., Tischler T.R., Cook A.G., Calleja N.D., Sloan T., Elliott D.A. & Sereno P. (2009) - New Mid-Cretaceous (Latest Albian) Dinosaurs from Winton, Queensland, Australia. PLoS ONE, 4(7): e6190.

Hone D. & Holtz T.R. Jr. (2017) - A century of spinosaurs - a review and revision of the Spinosauridae with comments on their ecology. Acta Geologica Sinica - English Edition. 91(3): 1120-32.

Hone D. & Holtz T.R. Jr. (2021) - Evaluating the ecology of Spinosaurus: shoreline generalist or aquatic pursuit specialist? Palaeontologia Electronica, 24(1): 1-28.

Ibrahim N., Sereno P.C., Dal Sasso C., Maganuco S., Fabbri M., Martill D.M., Zouhri S., Myhrvold N. & Iurino D.A. (2014) - Semiaquatic adaptations in a giant predatory dinosaur. Science, 345(6204): 1613-6.

Janensch W. (1955) - Der Ornithopode Dysalotosaurus der Tendaguru-Schichten. Palaeontographica, Supplement VII, 1(3): 105-176.

Janensch W. (1961) - Skelettrekonstruktion Von Dysalotosaurus Lettow-Vorbecki. Palaeontographica, Supplement VII, 3 (4): 237-240.

Kachhara R.P. & Jodhawat R.L. (1981) - On the age of Jaisalmer Formation, Rajasthan, India. Proceedings of IX Indian Colloquium on Micropalaeontology and Stratigraphy, Udaipur: 235-247.

Krishna J. (1979) - Callovian-Tithonian Ammonite Stratigraphy and Biogeography of Jaisalmer, India. Ammonoidea. Systematics Association Symposium, Abstracts: 33, York.

Madsen J. R. Jr. (1976) - Allosaurus fragilis: a revised osteology. Utah Geological Survey Bulletin, 109: 1-163.

Maganuco S., Cau A. & G. Pasini G. (2005) - First description of theropod remains from the Middle Jurassic (Bathonian) of Madagascar. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale in Milano, 146(II): 165-202.

Maganuco S., Cau A., Dal Sasso C. & Pasini G. (2007) - Evidence of large theropods from the Middle Jurassic of the Mahajanga Basin, NW Madagascar, with implications for ceratosaurian pedal ungual evolution. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale in Milano, 148: 261-271.

Maganuco S. & Dal Sasso C. (2018) - The smallest biggest theropod dinosaur: a tiny pedal ungual of a juvenile Spinosaurus from the Cretaceous of Morocco. PeerJ, 6: e4785.

Marsh O.C. (1879) - Notice of new Jurassic reptiles. American Journal of Science and Arts, 18(108): 501-505.

Marsh O.C. (1881) - Principal Characters of American Jurassic Dinosaurs, Part V. American Journal of Science, S3- 21(125): 417-23.

Mateus O. & Estraviz-López D. (2022) - A new theropod dinosaur from the Early Cretaceous (Barremian) of Cabo Espichel, Portugal: implications for spinosaurid evolution. PLoS ONE, 17: E0262614.

Mathur U.B., Pant S.C., Mehra S. & Mathur A.K. (1985) - Discovery of dinosaurian remains in Middle Jurassic of Jaisalmer, Rajasthan, Western India. Bulletin of the Indian Geologists’ Association, 18(2): 59-65.

Mcfeeters B., Ryan M.J. & Cullen T.M. (2018) - Positional variation in pedal unguals of North American ornithomimids (Dinosauria, Theropoda): a response to Brownstein (2017). Vertebrate Anatomy Morphology Palaeontology, 6: 60-67.

Moreno K., Carrano M. T. & Snyder R. (2007) - Morphological changes in pedal phalanges through ornithopod dinosaur evolution: a biomechanical approach. Journal of morphology, 268(1): 50-63.

Norman D. (1986) - On the anatomy of Iguanodon atherfieldensis (Omithischia: Omithopoda). Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, 56: 281-372.

Norman D.B., Witmer L.M. & Weishampel D.B. (2004) - Basal Thyreophora. In: Weishampel D.B., Dodson P., & Osmólska H. (Ed.) - The Dinosauria, 2nd Edition. University Of Californian Press: 335-342.

Novas F.E. & Bandyopadhyay S. (2001) - Abelisaurid pedal unguals from the Late Cretaceous of India. VII International Symposium on Mesozoic Terrestrial Ecosystems, Buenos Aires: 145-149.

Novas F.E., Cambiaso A.V. & Ambrioso A. (2004) - A new basal iguanodontian (Dinosauria, Ornithischia) from the Upper Cretaceous of Patagonia. Ameghiniana, 41(1): 75-82.

Novas F.E., Dalla Vecchia F.M. & Pais D. (2005) - Theropod pedal unguals from the Late Cretaceous (Cenomanian) of Morocco, Africa. MACN, F 7:167-75.

Oldham R.D. (1886) - Preliminary notes on the geology of Northern Jaisalmer. Records of the Geological Survey of India, 19(3): 157-160.

Ostrom J.H. (1970) - Stratigraphy and paleontology of the Cloverly Formation (Lower Cretaceous) of the Bighorn Basin area, Wyoming and Montana. Peabody Museum Bulletin, 35: 1-234.

Owen R. (1842) - Report on British fossil reptiles, part II. Report of the British Association for the Advancement of Science, 11: 60-294.

Owen R. (1869) - Description of a great part of a jaw with the teeth of Strophodus medius, from the Oolite of Caen in Normandy. Geological Magazine, 6: 193-196.

Pal T.K., Ray S.K., Talukder B. & Naik M.K. (2007) - On the mega-invertebrate fauna (Mollusca and Brachiopoda) of Cenozoic and Mesozoic of Jaisalmer, Rajasthan and their stratigraphic implications. Records of the Zoological Survey of India, Occasional Paper, 280: 1-40.

Pandey D.K., Fürsich F.T. & Alberti M. (2014) - Stratigraphy and Palaeoenvironments of the Jurassic Rocks of the Jaisalmer Basin - Field Guide. Beringeria, Special Issue 9: 1-111.

Pandey D.K., Fürsich F.T. & Baron-Szabo R. (2009) - Jurassic Corals from the Jaisalmer Basin, West Rajasthan, India. Zitteliana, A48/49: 13-37.

Parks W.A. (1926) - Thescelosaurus warreni, a new species of orthopodous dinosaur from the Edmonton Formation of Alberta. University of Toronto Studies, Geology Series, 21: 1-42.

Pieńkowski G., Brański P., Pandey D.K., Schlög J., Alberti M. & Fürsich F.T. (2015) - Dinosaur footprints from the Thaiat Ridge and their palaeoenvironmental background, Jaisalmer Basin, Rajasthan, India. Volumina Jurassica, 13(1): 17-26.

Posmoşanu E. (2003) - Iguanodontian dinosaurs from the lower Cretaceous bauxite site from Romania. Acta Palaeontologica Romaniae, 4: 431-439.

Prasad G.V.R. & Parmar. V. (2020) - First Ornithischian and theropod dinosaur teeth from the Middle Jurassic Kota Formation of India: paleobiogeographic relationships. In: Prasad G.V.R. & Patnaik R. (Eds.) - Biological consequences of plate tectonics: new perspectives on Post-Gondwana break-up - A tribute to Ashok Sahni. Vertebrate Paleobiology and Paleoanthropology Series: 1-30.

Rotatori F.M., Moreno-Azanza M. & Mateus O. (2022) - Reappraisal and new material of the holotype of Draconyx loureiroi (Ornithischia: Iguanodontia) provide insights on the tempo and modo of evolution of thumb-spiked dinosaurs. Zoological Journal of the Linnean Society, 195(1): 125-156.

Rozadilla S., Agnolín F.L. & Novas F.E. (2019) - Osteology of the Patagonian ornithopod Talenkauen santacrucensis (Dinosauria, Ornithischia). Journal of Systematic Palaeontology, 17(24): 2043-2089.

Sahni M.R. (1955) - Recent research in Palaeontologic Division, Geological Survey of India. Current Science, 24: 187.

Sahni M.R. & Bhatnagar N.C. (1958) - New fossils from the Jurassic rocks of Jaisalmer, Rajasthan. Records of the Geological Survey of India, 87(2): 428-437.

Sereno P.C. (2012) - Taxonomy, morphology, masticatory function and phylogeny of heterodontosaurid dinosaurs. Zookeys, 226: 1-225.

Sereno P.C., Beck A.L., Dutheil D.B., Gado B., Larsson H.C.E., Lyon G.H., Marcot J.D., Rauhut O.W.M., Sadleir R.W., Sidor C.A., Varricchio D.D., Wilson G.P. & Wilson J.A. (1998) - A long-snouted predatory dinosaur from Africa and the evolution of spinosaurids. Science, 282: 1298-1302.

Sereno P.C., Forster C.A., Larsson H.C.E., Dutheil D.B. & Sues H.D. (1994) - Early Cretaceous dinosaurs from the Sahara. Science, 266: 267-271.

Sereno P., Myhrvold N., Henderson D., Fish F., Vidal D., Baumgart S., Keillor T., Formoso K. & Conroy L. (2022) - Spinosaurus is not an aquatic dinosaur. Published online 2022 Nov 30. doi: 10.7554/eLife.80092 10.1101/2022.05.25.493395.

Serrano-Martínez A., Daniel V., Lara S., Francisco O. & Fabien K. (2016) - Isolated theropod teeth from the Middle Jurassic of Niger and the early dental evolution of Spinosauridae. Acta Palaeontologica Polonica, 61: 403-15.

Sharma A., Hendrickx C. & Singh S. (2023) - First Theropod record from the Marine Bathonian of Jaisalmer Basin, Tethyan Coast of Gondwanan India. Rivista Italiana di Paleontologia e Stratigrafia, 129(1): 49-64.

Sharma A. & Singh S. (2021) - A small assemblage of marine hybodont sharks from the Bathonian of the Jaisalmer Basin, India. Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen, 301(3): 317-333.

Sharma A., Singh S. & Satheesh S. R. (2022) - The first Turiasaurian sauropod of India reported from the Middle Jurassic (Bathonian) sediments of Jaisalmer Basin, Rajasthan, India. Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen, 304(2): 187-203.

Singh C.S.P., Jaitly A.K. & Pandey D.K. (1982) - First report of some Bajocian-Bathonian (Middle Jurassic) ammonoids and the age of the oldest sediments from Kachchh, Western India. Newsletters on Stratigraphy, 11: 37-40.

Singh C.S.P., Pandey D.K. & Jaitly A.K. (1983) - Discovery of Clydoniceras Blake and Gracilisphinctes Buckman (Bathonian-Middle Jurassic Ammonites) in Kachchh, Western India. Journal of Palaeontology, 57: 821-824.

Stromer (1915) - Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Bahariye-Stufe (unterstes Cenoman). 3. Das Original des Theropodes Spinosaurus aegyptiacus nov. Gen., nov. Spec. Abhandlungen der Königlich Bayerischen Akademie der Wissenschaften, Mathematisch-Physikalische Klasse, München 28: 1-28.

Stromer E. (1934) - Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens II. Wirbeltierreste der Baharije-Stufe (unterstes Cenoman) 13. Dinosauria Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung Neue Folge, Heft 22.

Virchow H. (1919) - Atlas und Epistropheus bei den Schildkröten. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 8: 303-332.

Weishampel D.B., Jianu C.M., Csiki Z. & Norman D.B. (2003) - Osteology and phylogeny of Zalmoxes (n. g.), an unusual euornithopod dinosaur from the Latest Cretaceous of Romania. Journal of Systematic Palaeontology, 1(2): 65-123.

Whitton M.P. (2015) - Were early pterosaurs inept terrestrial locomotors? PeerJ, 3. e1018. 10.7717/peerj.1018.

Yadagiri P. (1988) - A new sauropod Kotasaurus yamanpalliensis from Lower Jurassic Kota Formation of India. Records of the Geological Survey of India, 11: 102-127.

Yadagiri p. (2001) - The osteology of Kotasaurus yamanpalliensis, a sauropod dinosaur from the Early Jurassic Kota Formation of India. Journal of Vertebrate Paleontology, 21 (2): 242 252.

Zaher H., Pol D., Navarro B.A., Delcourt R. & Carvalho A.B. (2020) - An Early Cretaceous theropod dinosaur from Brazil sheds light on the cranial evolution of the Abelisauridae. Comptes Rendus Palevol, 19(6): 101-115.






Received 2023-04-22
Accepted 2023-09-22
Published 2023-10-27