A New Troodontid Dinosaur from the Lower Cretaceous Yixian Formation of Liaoning Province, China
Corresponding Author
Caizhi SHEN
Institute of Geology, Chinese Academy of Geological Sciences, Beijing 100037 China
Dalian Natural History Museum, Dalian, Liaoning 116023 China
Corresponding author. E-mail: [email protected]Search for more papers by this authorJunchang LÜ
Institute of Geology, Chinese Academy of Geological Sciences, Beijing 100037 China
Search for more papers by this authorSizhao LIU
Dalian Natural History Museum, Dalian, Liaoning 116023 China
Search for more papers by this authorMartin KUNDRÁT
Department of Ecology, Faculty of Natural Sciences, Comenius University, Bratislava, Slovak Republic
Center for Interdisciplinary Biosciences, Faculty of Science, University of Pavol Jozef Šafárik, Košice, Slovak Republic
Search for more papers by this authorStephen L. BRUSATTE
University of Edinburgh, School of GeoSciences, Edinburgh EH4 2BH, UK
Search for more papers by this authorHailong GAO
Institute of Geomechanics, Chinese Academy of Geological Sciences, Beijing 100037 China
Search for more papers by this authorCorresponding Author
Caizhi SHEN
Institute of Geology, Chinese Academy of Geological Sciences, Beijing 100037 China
Dalian Natural History Museum, Dalian, Liaoning 116023 China
Corresponding author. E-mail: [email protected]Search for more papers by this authorJunchang LÜ
Institute of Geology, Chinese Academy of Geological Sciences, Beijing 100037 China
Search for more papers by this authorSizhao LIU
Dalian Natural History Museum, Dalian, Liaoning 116023 China
Search for more papers by this authorMartin KUNDRÁT
Department of Ecology, Faculty of Natural Sciences, Comenius University, Bratislava, Slovak Republic
Center for Interdisciplinary Biosciences, Faculty of Science, University of Pavol Jozef Šafárik, Košice, Slovak Republic
Search for more papers by this authorStephen L. BRUSATTE
University of Edinburgh, School of GeoSciences, Edinburgh EH4 2BH, UK
Search for more papers by this authorHailong GAO
Institute of Geomechanics, Chinese Academy of Geological Sciences, Beijing 100037 China
Search for more papers by this authorAbout the first author:
SHEN Caizhi, male, born in 1977; a Ph.D student at Institute of Geology, Chinese Academy of Geological Sicences; major in stratigraphy and palaeotoglogy;.
Email: [email protected]
Abstract
A new troodontid dinosaur, Daliansaurus liaoningensis gen. et sp. nov., is erected based on a nearly complete specimen from the Lower Cretaceous Yixian Formation of Beipiao City, Liaoning Province, China. This well preserved skeleton provides important new details of the anatomy for Liaoning troodontids, and gives new insight into their phylogenetic relationships and evolution. Daliansaurus is distinguished from other troodontids by an enlarged ungual on pedal digit IV, which is approximately the same size as the sickle-shaped second ungual, and is differentiated from other Liaoning troodontids by a number of characters of the skull, manus, pelvis, and hindlimb. A phylogenetic analysis recovers Daliansaurus within a subclade of Liaoning troodontids that also includes Sinovenator, Sinusonasus, and Mei. We erect a name for this group—Sinovenatorinae—and argue that it reflects a localized radiation of small-bodied troodontids in the Early Cretaceous of eastern Asia, similar to previously recognized radiations of Liaoning dromaeosaurids and avialans. As more Liaoning theropods are discovered, it is becoming apparent that small, feathered paravians were particularly diverse during the Early Cretaceous, and future work is needed to clarify how this diversity arose, which species coexisted, and how these numerous species partitioned niches.
References
- Agnolin, F.L, and Novas, F.E., 2013. Avian Ancestors. Herdelberg: Springer.
10.1007/978-94-007-5637-3 Google Scholar
- Averianov, A.O. and Sues, H.D., 2007. A new troodontid (Dinosauria: Theropoda) from the Cenomanian of Uzbekistan, with a review of troodontid records from the territories of the Former Soviet Union. Journal of Vertebrate Paleontology, 27: 87–98.
- Barsbold, R., 1974. Saurornithoididae, a new family of carnivorous dinosaurs from Central Asia and North America. Palaeontologia Polonica, 30: 5–22.
- Barsbold, R., Osmólska, H., and Kurzanov, S.M., 1987. On a new troodontid (Dinosauria, Theropoda) from the Early Cretaceous of Mongolia. Acta Palaeontologica Polonica, 32: 121–132.
- Barsbold, R., and Osmólska, H., 1999. The skull of Velociraptor from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica, 44: 189–219.
- Brusatte, S.L., Lloyd, G.T., Wang, S.C., and Norell, M.A., 2014. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Current Biology, 24: 2386–2392.
- Burea of Geology and Mineral Resources of Liaoning Province., 1989. Regional geology of Liaonig Province. Beijing: Geological Publishing House, 690–691 (in Chinese).
- Cau, A., Brougham, T., and Naish, D., 2015. The phylogenetic affinities of the bizarre Late Cretaceous Romanian theropod Balaur bondoc (Dinosauria, Maniraptora): dromaeosaurid or flightless bird? PeerJ, Doi: 10.7717/peerj.1032
- Chatterjee, S., 2015. The rise of birds: 225 million years of evolution. Johns Hopkins University Press, 45–48.
- Chiappe, L.M., Norell, M.A., and Clark, J.M., 1998. The skull of a new relative of the stem-group bird Mononykus. Nature, 392: 275–278.
- Codd, J.R., Manning, P.L., Norell, M.A., and Perry, S. F., 2008. Avian-like breathing mechanics in maniraptoran dinosaurs. Proceedings of the Royal Society, Series B 275: 157–161.
- Cood, J.R., 2010. Uncinate process in birds: morphology, physiology and function. Comparative Biochemistry and Physiology, Part A 156, 303–308.
- Currie, P.J., and Dong, Z.M., 2001. New information on Cretaceous troodontids (Dinosauria, Theropoda) from the People's Republic of China. Canadian Journal of Earth Sciences, 38: 1753–1766.
- Currie, P.J., 1987. Bird-like characteristics of the jaws and teeth of troodontid theropods (Dinosauria, Saurischia). Journal of Vertebrate Paleontology, 7: 72–81.
10.1080/02724634.1987.10011638 Google Scholar
- Gauthier, J., 1986. Saurischian monophyly and the origin of birds. In: K. Padian (ed.), The origin of birds and the evolution of flight. Memoirs of the California Academy of Sciences, 8: 1–55.
- Gao, C.L., Morschhauser, E.M., Varricchio, D.J., Liu, J.Y., and Zhao, B., 2012. A second soundly sleeping dragon: new anatomical details of the Chinese troodontid Mei long with implications for phylogeny and taphonomy. PLoS One, 7: e45203.
- Gilmore, C.W., 1924. On Troodon validus, an ornithopodous dinosaur from the Belly River Cretaceous of Alberta, Canada. Bulletin of Department of Geology, University of Alberta 1: 1–143.
- Godefroit, P., Cau, A., Hu, D.Y., Escuillié, F., Wu, W.H., and Dyke, G., 2013a. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature, 498: 359–362.
- Godefroit, P., Demuynck, H., Dyke, G., Hu, D.Y., Escuillié, F., and Claeys, P., 2013b. Reduced plumage and flight ability of a new Jurassic paravian theropod from China. Nature Communications, 4: 1394.
- Goswami, A., Prasad, G.V.R., Verma, O., Flynn, J.J., and Benson, R.B.J, 2013. A troodontid dinosaur from the latest Cretaceous of India. Nature Communications, 4: 1703.
- Han, G., Chiappe, L.M., Ji, S.A., Habib, M., Turner, A.H., Chinsamy, A., Liu, X.L and Han, L.Z. 2014. A new raptorial dinosaur with exceptionally long feathering provides insights into dromaeosaurid flight performance. Nature Communications. 5: 4382.
- He, Z.J., Niu, B.G., and Zhang, X.Y., 2007. Later Jurassic thrusting and syntectonic sedimentary basin systems in the Chaoyang Region, Western Liaoning. Geological Review, 53: 152–163 (in Chinese).
- Hu, D.Y., Hou, L.H., Zhang, L.J., and Xu, X., 2009. A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus. Nature, 461: 640–643.
- Ji, Q., Norell, M.A., Makovicky, P.J., Gao, K.Q. Ji S.A., and Yuan C.X., 2003. An early ostrich dinosaur and implications for ornithomimosaur phylogeny. American Museum Novitates, 3420: 1–19.
- Ji, Q., Ji, S.A., Lu, J.C., You, H.L., Chen, W., Liu, Y.Q., and Liu, Y.X., 2005. First avialian bird from China. Geological Bulletin of China, 24: 197–210.
- Ji, S.A., and Ji, Q., 2007. Jinfengopteryx Compared to Archaeopteryx, with comments on the Mosaic evolution of long-tailed Avialan birds. Acta Geologica Sinica (English Edition), 81: 337–343.
- Kurzanov, S.M., and Osmólska, H., 1991. Tochisaurus nemegtensis gen. et sp. n., a new troodontid (Dinosauria, Theropoda) from Mongolia. Acta Palaeontologica Polonica, 36: 69–76.
- Lü, J.C., 2002. A new Oviraptorosaurid (Theropoda: Oviraptorosauria) from the Late Cretaceous of Southern China. Journal of Vertebrate Paleontology, 22: 871–875.
- Lü, J.C., Xu, L., Liu, Y.Q., Zhang, X.L., Jia, S.H., and Ji, Q., 2010. A new troodontid theropod from the Late Cretaceous of Central China, and the radiation of Asian Troodontids. Acta Palaeontologica Polonica, 55: 381–388.
- Lü, J.C., and Brusatte, S.L., 2015. A large, short-armed, winged dromaeosaurid (Dinosauria: Theropoda) from the Early Cretaceous of China and its implications for feather evolution. Science Report, 5: 11775.
- Makovicky, P.J., Norell, M.A., Clark, J.M., and Rowe, T., 2003. Osteology and relationships of Byronosaurus jaffei (Theropoda: Troodontidae). American Museum Novitates, 3402: 1–32.
- Makovicky, P.J., and Norell, M.A., 2004. Troodontidae. In: D.B. Weishampel, P. Dodson, and H. Osmóska, (eds), The Dinosauria. Second edition. Berkeley: University of California Press, 184–195.
10.1525/california/9780520242098.003.0011 Google Scholar
- Marsh, O.C., 1881. Classification of the Dinosauria. American Journal of Science (series 3), 23: 81–86.
- Norell, M.A., and Makovicky, P.J., 1999. Important features of the dromaeosaurid skeleton II: information from newly collected specimens of Velociraptor mongoliensis. American Museum Novitates, 3282: 1–45.
- Norell, M.A., Makovicky, P.J., and Clark, J.M., 2000. A new troodontid theropod from Ukhaa Tolgod, Mongolia. Journal of Vertebrate Paleontology, 20: 1–11.
- Norell, M.A., Clark, J.M., and Makovicky, P.J., 2001. Relationships among Maniraptora: problems and prospects. In: J. Gauthier, and L.F. Gall (eds.), New Perspectives on the Origin and Early Evolution of Birds. New Haven: Peabody Museum of Natural History, 49–67.
- Norell, M.A., Makovicky, P.J., Bever, G.S., Balanoff, A.M., Clark, J.M., Barsbold, R., and Rowe, T., 2009. A review of the Mongolian Cretaceous dinosauru Saurornithoides (Troodontidae: Theropoda). American Museum Novitates, 3654: 1–63.
10.1206/648.1 Google Scholar
- Osmólska, H., 1987. Borogovia Gracilicrus gen.et sp. n., A new troodontid dinosaur from the Late Cretaceous of Mongolia. Palaeontologica, 32: 133–150.
- Ostrom, J.H. 1969a. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History, 30: 1–165.
- Rowe, T., 1986. Homology and evolution of the deep dorsal thigh musculature in birds and other reptilia. Journal of Morphology, 189: 327–346.
- Rueesll, D.A., 1948. The dentary of Troodon, a genus of Theropod Dinosaurs. Journal of Paleontology, 22: 625–629.
- Russell, D.A., 1969. A new specimen of Stenonychosaurus from the Oldman Formation (Cretaceous) of Alberta. Canadian Journal of Earth Sciences, 6: 595–612.
- Russell, D.A., and Dong, Z.M., 1993. A nearly complete skeleton of a new troodontid dinosaur from the Early Cretaceous of the Ordos Basin, Inner Mongolia, People's Republic of China. Canadian Journal of Earth Sciences, 30: 2163–2173.
- Senter, P., Kirkland, J.I., DeBlieux, D.D., Madsen, S., and Toth, N., 2012. New Dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the Evolution of the Dromaeosaurid Tail. PLoS One, 7: e36790.
- Tsuihiji, T., Barsbold, R., Watabe, M., Tsogtbaatar, K., Chinzorig, T., Fujiyama, Y., and Suzuki, S., 2014. An exquisitely preserved troodontid theropod with new information on the palatal structure from the Upper Cretaceous of Mongolia. Naturwissenschaften, 101: 131–142.
- Tsuihiji, T., Barsbold, R., Watabe, M., Tsogtbaatar, K., Suzuki, S., and Hattori, S., 2016. New material of a troodontid theropod (Dinosauria: Saurischia) from the Lower Cretaceous of Mongolia. Historical Biology, 28: 128–138.
- Turner, A.H., Makovicky, P.J., and Norell, M.A., 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History, 371: 1–206.
10.1206/748.1 Google Scholar
- Xu, X., Wang, X.L., and Wu, X.C., 1999. A dromaeosaurid dinosaur with a fliamentous integument from the Yixian Formation of China. Nature, 401: 262–266
- Xu, X., Zhou, Z.H., and Wang, X.L., 2000. The smallest known non-avian theropod dinosaur. Nature, 408: 705–708.
- Xu, X., 2002. Deinonychosaurian fossils from the Jehol Group of western Liaoning and the coelurosaurian evolution. Beijing: Chinese Academy of Sciences, (Ph. D thesis): 322 p.
- Xu, X., Norell, M.A., Wang, X.L., Makovicky, P.J., and Wu, X.C., 2002. A basal troodontid from the Early Cretaceous of China. Nature, 415: 780–784.
- Xu, X., Zhou, Z.H., Wang, X.L., Kuang, X.W., Zhang, F.C., and Du, X.K., 2003. Four-winged dinosaurs from China. Nature, 421: 335–340.
- Xu, X., and Norell, M.A., 2006. Non-Avian dinosaur fossils from the Lower Cretaceous Jehol Group of western Liaoning, China. Geological Journal, 41: 419–437.
- Xu, X., and Wang, X.L., 2004. A new troodontid (Theropoda: Troodontidae) from the Lower Cretaceous Yixian Formation of Western Liaoning, China. Acta Geologica Sinica (English Edition), 78(1): 22–26.
- Xu, X., and Norell, M.A., 2004. A new troodontid dinosaur from China with avian-like sleeping posture. Nature, 431: 838–841.
- Xu, X., Tan, J.W., Wang, J.M., Zhao, X.J., and Tan, L., 2007. A gigantic bird-like dinosaur from the Late Cretaceous of China. Nature, 447: 844–847.
- Xu, X., Zhao, Q., Norell, M.A., Sullivan, C., Hone, D., Erickson, G., Wang, X.L., Han, F.L., and Guo, Y., 2008. A new feathered maniraptoran dinosaur fossil that fills a morphological gap in avian origin. Chinese science bulletin, 54: 430–435.
- Xu, X., Choiniere, J.N., Pittman, M., Tan, Q.W., Xiao, D., Li, Z.Q., Tan, L., Clark, J. M., Norell, M.A., Hone, D.W.E, and Sullivan, C., 2010. A new dromaeosaurid (Dinosauria: Theropoda) from the Upper Cretaceous Wulansuhai Formation of Inner Mongolia, China. Zootaxa, 2403: 1–9.
10.11646/zootaxa.2403.1.1 Google Scholar
- Xu, X., Tan, Q.W., Sullivan, C., Han, F.L., and Xiao, D., 2011a. A short-armed troodontid dinosaur from the Upper Cretaceous of Inner Mongolia and its implications for troodontid evolution. PloS One, 6: e22916.
- Xu, X., You, H.L., Du, K., and Han, F.L., 2011b. An Archaeopteryx-like theropod from China and the origin of Avialae. Nature, 475: 465–470.
- Xu, X., Zhao, Q., Sullivan, C., Tan, Q.W., Sander, M., and Ma, Q.Y., 2012. The taxonomy of the troodontid IVPP V 10597 reconsidered. Vertebr PalAsiat, 50: 140–150.
- Xu, X., Zhou, Z.H., Dudley, R., Mackem, S., Chuong, C.M., Erickson, G.M., and Varricchio, D.J., 2014. An integrative approach to understanding bird origins. Science, 346.
- Zanno, L.E., Varricchio, D.J., O'Connor, P.M., Titus, A.L., and Knell, M.J., 2011. A new troodontid theropod, Talos sampsoni gen. et sp. nov., from the Upper Cretaceous Western Interior Basin of North America. PLoS One, 6: e24487.
- Zhang, H., Liu, X.M., Li, Z.T., Yang, F.L., and Wang, X.R., 2005. Early Cretaceous large-scale crustal thinning in the Fuxin-Yixian basin and adjacent area in Western Liaoning. Geological Review, 51: 360–372 (in Chinese).
- Zheng, X.H., Xu, X., You, H.L., Zhao, Q., Dong, Z.M., 2010. A short-armed dromaeosaurid from the Jehol Group of China with implications for early dromaeosaurid evolution. Proceedings of the Royal Society B. 277 (1679): 211–217.