Abstract
Hungarosaurus tormai is a medium-sized nodosaurid ankylosaur that was described on the basis of four partial skeletons from the Santonian (Upper Cretaceous) of Hungary. In this paper, a new fifth skeleton and several isolated remains are described which greatly improve our knowledge of this primitive nodosaurid ankylosaur. Isolated cranial remains referred to juvenile individuals provide new information on the development of cranial ornamentation in nodosaurid ankylosaurs. Apart from both preserved mandibles with in situ dentition, the fifth partial skeleton contains several previously unknown limb elements (humerus, ulnae, radius) that indicate unusual limb proportions for Hungarosaurus compared with other ankylosaurs. On the basis of the five partial skeletons and the isolated remains, a skeletal and dermal armor reconstruction is attempted. Body mass calculations using three different methods yield an estimate of 650 kg for H. tormai.
Kurzfassung
Hungarosaurus tormai ist ein mittelgroßer Nodosauride, der ursprünglich auf der Basis von vier Teilskeletten aus dem Santon (Oberkreide) von Ungarn beschrieben wurde. In dieser Arbeit wird ein fünftes Skelett und einige isolierte Elemente beschrieben, die unsere Kenntnis dieses primitiven Nodosauriden deutlich verbessern. Isolierte Schädelelemente von Jungtieren liefern neue Informationen zur Entwicklung der Schädelornamentation bei Nodosauriden. Neben beiden erhaltenen Unterkiefern, zum Teil noch mit Zähnen in Situ, enthält das fünfte Skelett einige bisher nicht bekannte Extremitätenknochen (Humerus, Ulnae, Radius), die darauf hinweisen, daß Hungarosaurus im Vergleich mit anderen Ankylosaurirn ungewöhnliche Bein-Proportionen besaß. Anhand der fünf Teilskelette und der isolierten Reste wird eine Rekonstruktion des gesamten Skelettes und der Panzerung versucht. Drei verschiedene Methoden der Rekonstruktion der Körpermasse ergaben eine Schätzung von 650 kg für H. tormai.
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References
Brown, B. 1908. The Ankylosauridae, a new family of armored dinosaurs from the Upper Cretaceous. Bulletin of the American Museum of Natural History 24: 187–201.
Bunzel, E. 1871. Die Reptilfauna der Gosau Formation in der neuen Welt bei Wiener—Neustadt. Abhandlungen der Kaiserliche und Königliche Geologische Reichsanstalt 5: 1–18.
Carpenter, K. 1982. Skeletal and dermal armor reconstruction of Euoplocephalus tutus (Ornithischia: Ankylosauridae) from the Late Cretaceous Oldman Formation of Alberta. Canadian Journal of Earth Sciences 19: 689–697.
Carpenter, K. 1984. Skeletal reconstruction and life restoration of Sauropelta (Ankylosauria: Nodosauridae) from the Cretaceous of North America. Canadian Journal of Earth Sciences 21: 1491–1498.
Carpenter, K. 1990. Ankylosaur systematics: example using Panoplosaurus and Edmontonia (Ankylosauria: Nodosauridae). In Dinosaur systematics: Approaches and perspectives, ed. K. Carpenter, and P. Currie, 281–298. Cambridge: Cambridge University Press.
Carpenter, K. 1997. Ankylosaurs. In The complete sinosaur, ed. J.O. Farlow, and M.K. Brett-Surman, 307–316. Indianapolis: Indiana University Press.
Carpenter, K. 2004. Redescription of Ankylosaurus magniventris Brown, 1908 (Ankylosauridae) from the Upper Cretaceous of the Western Interior of North America. Canadian Journal of Earth Sciences 41: 961–986.
Carpenter, K., D. Dilkes, and D.B. Weishampel. 1995. The dinosaurs of the Niobrara Chalk Formation (Upper Cretaceous, Kansas). Journal of Vertebrate Paleontology 15: 275–297.
Carpenter, K., and J.I. Kirkland. 1998. Review of Lower and Middle Cretaceous Ankylosaurs from North America. In New Mexico museum of natural history and science bulletin 14, ed. S. Lucas, J.I. Kirkland, and J.W. Estep, 249–270. Albuquerque: Lower and Middle Cretaceous Terrestrial Ecosystems.
Colbert, E.C. 1962. The weights of dinosaurs. American Museum Novitates 2076: 1–16.
Coombs, P.W. 1978. Forelimb muscles of the Ankylosauria (Reptilia, Ornithischia). Journal of Paleontology 52: 642–657.
Coombs, P.W. 1979. Osteology and myology of the hindlimb in the Ankylosauria (Reptilia, Ornithischia). Journal of Paleontology 53: 666–684.
Coombs, P.W. 1990. Teeth and taxonomy in ankylosaurs. In Dinosaur systematics: Approaches and perspectives, ed. K. Carpenter, and P.J. Currie, 269–279. Cambridge: Cambridge University Press.
Coombs, P.W. 1995. A new nodosaurid ankylosaur (Dinosauria: Ornithischia) from the Lower Cretaceous of Texas. Journal of Vertebrate Paleontology 15: 298–312.
Coombs, P.W., and T. Maryanska. 1990. Ankylosauria. In The Dinosauria, ed. D.B. Weishampel, P. Dodson, and H. Osmólska, 456–483. Berkeley: California University Press.
Eaton, T.H. 1960. A new armored dinosaur from the Cretaceous of Kansas. The University of Kansas Paleontological Contributions 25: 1–24.
Ford, T.L., and J.I. Kirkland. 2001. Carlsbad ankylosaur (Ornithischia, Ankylosauria): an ankylosaurid and not a nodosaurid. In The Armored Dinosaurs, ed. K. Carpenter, 239–260. Bloomington: Indiana University Press.
Galton, P.M. 1983. Armored dinosaurs (Ornithischia: Ankylosauria) from the Middle and Upper Jurassic of Europe. Palaeontographica (Abteilung A) 182: 1–25.
Garcia, G., and X. Pereda-Suberbiola. 2003. A new species of Struthiosaurus (Dinosauria: Ankylosauria) from the Upper Cretaceous of Villeveyrac (southern France). Journal of Vertebrate Paleontology 23: 156–165.
Gaston, R.W., J. Schellenbach, and J.I. Kirkland. 2001. Mounted skeleton of the polacanthine ankylosaur Gastonia burgei. In The armored dinosaurs, ed. K. Carpenter, 386–398. Bloomington: Indiana University Press.
Haas, G. 1969. On the jaw muscles of ankylosaurs. American Museum Novitates 2399: 1–11.
Hill, R.W., L.M. Witmer, and M.A. Norell. 2003. A new specimen of Pinacosaurus grangeri (Dinosauria: Ornithischia) from the Late Cretaceous of Mongolia: ontogeny and phylogeny of ankylosaurs. American Museum Novitates 3395: 1–29.
Henderson, D. 1999. Estimating the masses and centers of mass of extinct animals by 3D mathematical slicing. Paleobiology 25: 88–106.
Hulke, J.W. 1887. Supplementary note on Polacanthus foxii, describing the dorsal shield and some parts of the endoskeleton, imperfectly known in 1881. Philosophical Transactions of the Royal Society of London 178: 169–172.
Huxley, T.H. 1867. On Acanthopholis horridus, a new reptile from the Chalk Marl. Geological Magazine 4: 65–67.
Jocha-Edelényi, E. 1988. History of evolution of the Upper Cretaceous basin in the Bakony Mts at the time of the terrestrial Csehbánya Formation. Acta Geologica Hungarica 31: 19–31.
Kirkland, J.I. 1998. A polacanthine ankylosaur (Ornithischia: Dinosauria) from the Early Cretaceous (Barremian) of Eastern Utah. In New Mexico museum of natural history and science bulletin 14, ed. S. Lucas, J.I. Kirkland, and J.W. Estep, 271–281. Albuquerque: Lower and Middle Cretaceous Terrestrial Ecosystems.
Knauer, J., and Á. Siegl-Farkas. 1992. Palynostratigraphic position of the Senonian beds overlying the Upper Cretaceous bauxite formations of the Bakony Mts. Annual Report of the Hungarian Geological Institute of 1990, pp 463–471
Lee, J.-N. 1996. A new nodosaurid ankylosaur (Dinosauria: Ornithischia) from the Paw Paw Formation (Late Albian) of Texas. Journal of Vertebrate Paleontology 16: 232–245.
Lull, R.S. 1921. The Cretaceous armored dinosaur, Nodosaurus textilis Marsh. The American Journal of Science (5. Series) 1: 97–132.
Marsh, C.O. 1890. Additional characters of the Ceratopsidae with notice of new Cretaceous dinosaurus. The American Journal of Science (3. Series) 39: 418–426.
Maryanska, T. 1977. Results of the Polish-Mongolian palaeontological expeditions-part VII: Ankylosauridae (Dinosauria) from Mongolia. Palaeontologia Polonica 37: 85–151.
Márton-Szalai, E. 2005. Paleomagnetic studies at Iharkút. Unpublished manuscript, ELTE Department of Applied Geology, Budapest
Molnar, R.E. 1980. An Ankylosaur (Ornithischia: Reptilia) from the Lower Cretaceous of southern Queensland. Memoirs of the Queensland Museum 20: 77–87.
Molnar, R.E. 2001. Armor of the small ankylosaur Minmi. In The armored dinosaurs, ed. K. Carpenter, 341–362. Bloomington: Indiana University Press.
Molnar, R.E., and E. Frey. 1987. The paravertebral elements of the Australian ankylosaur Minmi (Reptilia: Ornithischia, Cretaceous). Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 175: 19–37.
Nopcsa, F. 1915. Die Dinosaurier der siebenbürgischen Landesteile Ungarns. Mitteilungen des Jahrbuchs der Königliche Ungarische Geologische Reichsanstalt 23: 1–26.
Nopcsa, F. 1929. Dinosaurierreste aus Siebenbürgen V. Geologica Hungarica series Palaeontologica 4: 1–76.
Norman, D.B. 1984. A systematic reappraisal of the reptile order Ornithischia. In Third symposium on mesozoic terrestrial ecosystems, short papers, ed. W.-E. Reif, and F. Westphal, 157–162. Tübingen: Attempto Verlag.
Osborn, H.F. 1923. Two Lower Cretaceous dinosaurs from Mongolia. American Museum Novitates 95: 110.
Ostrom, J. 1970. Stratigraphy and paleontology of the Cloverly Formation (Lower Cretaceous) of the Bighorn Basin area, Wyoming and Montana. Bulletin of the Peabody Museum of Natural History 35: 1–134.
Ősi, A. 2004. The first dinosaur remains from the Upper Cretaceous of Hungary (Csehbánya Formation, Bakony Mts). Geobios 37: 749–753.
Ősi, A. 2005. Hungarosaurus tormai, a new ankylosaur (Dinosauria) from the Upper Cretaceous of Hungary. Journal of Vertebrate Paleontology 25: 370–383.
Pereda-Suberbiola, X. 1992. A revised census of European Late Cretaceous nodosaurids (Ornithischia: Ankylosauria): last occurrence and possible extinction scenarios. Terra Nova 4: 641–648.
Pereda-Suberbiola, X. 1999. Ankylosaurian dinosaur remains from the Upper Cretaceous of Laño (Iberian Peninsula). Estudios del Museo de Ciencias Naturales de Alava 14: 273–288.
Pereda-Suberbiola, X., and P.M. Barrett. 1999. A systematic review of ankylosaurian dinosaur remains from the Albian-Cenomanian of England. Special papers in Palaeontology 60: 177–208.
Pereda-Suberbiola, X., and P. Galton. 2001. Reappraisal of the nodosaurid ankylosaur Struthiosaurus austriacus Bunzel from the Upper Cretaceous Gosau beds of Austria. In The armored dinosaurs, ed. K. Carpenter, 173–210. Bloomington: Indiana University Press.
Rybczynski, N., and M.K. Vickaryous. 2001. Evidence of complex jaw movement in the Late Cretaceous ankylosaurid Euoplocephalus tutus (Dinosauria: Thyreophora). In The armored dinosaurs, ed. K. Carpenter, 299–317. Bloomington: Indiana University Press.
Seebacher, F. 2001. A new method to calculate allometric length–mass relationships of dinosaurs. Journal of Vertebrate Paleontology 21: 51–60.
Seeley, H.G. 1881. The reptile fauna of the Gosau Formation preserved in the Geological Museum of the University of Vienna. Quarterly Journal of the Geological Society London 37: 620–702.
Seeley, H.G. 1888. The classification of the Dinosauria. Report of the British Association of Advancement of Science 1887: 698–699.
Sereno, P.C., H.C.E. Larsson, C.A. Sidor, and B. Gado. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516–1519.
Swofford, D.L. 2000. PAUP: phylogenetic analysis using parsimony, version 4.0b10. Sunderland: Sinauer Associates.
Tuba, G.Y., P. Kiss, M. Pósfai, and A. Mindszenty. 2006. Preliminary data on the diagenesis of Cretaceous dinosaur bones from the Bakony Mts, Hungary. Földtani Közlöny 136: 1–24. (In Hungarian).
Vickaryous, M.K., A.P. Russell, and P.J. Currie. 2001. Cranial ornamentation of ankylosaurs (Ornithischia: Thyreophora): reappraisal for developmental hypotheses. In The armored dinosaurs, ed. K. Carpenter, 318–340. Bloomington: Indiana University Press.
Vickaryous, M.K., T. Maryanska, and D.B. Weishampel. 2004. Ankylosauria. In The Dinosauria, 2nd ed, ed. D.B. Weishampel, P. Dodson, and H. Osmólska, 363–392. Berkeley: The University of California Press.
Xu, X., X.-L. Wang, and H.-L. You. 2001. A juvenile ankylosaur from China. Naturwissenschaften 88: 297–300.
Acknowledgments
We are especially grateful to S.C.R. Maidment and two anonymous reviewers for making significant improvements to the manuscript. We thank the 2000–2006 field crews for their assistance in the field work and the Bakony Bauxite Mining Company for logistical help. We wish to thank P. Gulyás, T. Aranyi, Z. Szentesi, and A. Nyerges for preparation of the fifth skeleton. We are especially grateful to F. Szőke for producing a 3D frame for body mass estimation and to M. Vickaryous (Dalhousie University, Halifax, USA) for discussions. We thank N. Vávra (PIUW), C. Corral (MCNA), X. Pereda-Suberbiola (Universidad del País Vasco, Bilbao, Spain), I. Rutzky, J. Galkin, and C. Mehling (AMNH, New York, USA), D.B. Weishampel (Johns Hopkins University, Baltimore, USA), M.K. Brett-Surman (USNM, Washington DC, USA), and K. Carpenter (DMNH, Denver, USA) for access to material in their care. We are grateful to Massimo Delfino and Paolo Piras for their help in calculating decay indices. We thank J. Pálfy and S.C.R. Maidment for reading and correcting the earlier versions of the manuscript and for their useful suggestions. Fieldwork was financially supported by the Hungarian Natural History Museum, the National Geographic Society (Grant No. 7228-02, 7508-03), the Jurassic Foundation, the Hungarian Scientific Research Fund (Grant No. OTKA T38045), the Hantken Foundation, and the Hungarian Oil and Gas Company (MOL). This is an MTA–MTM Paleo contribution, no. 67.
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Appendix 1
Appendix 1
Data matrix used in the phylogenetic analysis corresponds to the data matrix used by Ősi (2005) with the additional information and modifications provided below:
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Ősi, A., Makádi, L. New remains of Hungarosaurus tormai (Ankylosauria, Dinosauria) from the Upper Cretaceous of Hungary: skeletal reconstruction and body mass estimation. Paläontol Z 83, 227–245 (2009). https://doi.org/10.1007/s12542-009-0017-5
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DOI: https://doi.org/10.1007/s12542-009-0017-5