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New remains of Hungarosaurus tormai (Ankylosauria, Dinosauria) from the Upper Cretaceous of Hungary: skeletal reconstruction and body mass estimation

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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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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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Correspondence to Attila Ősi.

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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