Abstract
The early evolution of the major groups of derived non-avialan theropods is still not well understood, mainly because of their poor fossil record in the Jurassic. A well-known result of this problem is the ‘temporal paradox’ argument that is sometimes made against the theropod hypothesis of avian origins1. Here we report on an exceptionally well-preserved small theropod specimen collected from the earliest Late Jurassic Tiaojishan Formation of western Liaoning, China2. The specimen is referable to the Troodontidae, which are among the theropods most closely related to birds. This new find refutes the ‘temporal paradox’1 and provides significant information on the temporal framework of theropod divergence. Furthermore, the extensive feathering of this specimen, particularly the attachment of long pennaceous feathers to the pes, sheds new light on the early evolution of feathers and demonstrates the complex distribution of skeletal and integumentary features close to the dinosaur–bird transition.
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References
Feduccia, A. Birds are dinosaurs: simple answer to a complex problem. Auk 119, 1187–1201 (2002)
Xu, K. et al. Jurassic System in the North of China (VII): The Stratigraphic Region of Northeast China (Petroleum Industry Press, 2003)
Xu, X. et al. A new feathered maniraptoran dinosaur fossil that fills a morphological gap in avian origin. Chin. Sci. Bull. 54, 430–435 (2009)
Wellnhofer, P. Archaeopteryx—Der urvogel von Solnhofen (Friedrich Pfeil, 2008)
Xu, X. & Norell, M. A. A new troodontid from China with avian-like sleeping posture. Nature 431, 838–841 (2004)
Norell, M. A. & Makovicky, P. J. in The Dinosauria 2nd edn (eds Weishampel, D. B., Dodson, P. & Osmolska, H.) 196–209 (Univ. California Press, 2004)
Makovicky, P. J. & Norell, M. A. in The Dinosauria 2nd edn (eds Weishampel, D. B., Dodson, P., Osmolska, H.) 184–195 (Univ. California Press, 2004)
Makovicky, P. J., Norell, M. A., Clark, J. M. & Rowe, T. Osteology and relationships of Byronosaurus jaffei (Theropda: Troodontidae). Am. Mus. Novit. 3402, 1–32 (2003)
Currie, P. J. Bird-like characteristics of the jaws and teeth of troodontid theropods (Dinosauria: Saurischia). J. Vert. Paleontol. 7, 72–81 (1987)
Makovicky, P. J. Apesteguía, S. & Agnolín, F. L. The earliest dromaeosaurid theropod from South America. Nature 437, 1007–1011 (2005)
Xu, X. & Wang, X.-L. A new dromaeosaur (Dinosauria: Theropoda) from the Early Cretaceous Yixian Formation of western Liaoning. Vert. PalAsia. 42, 111–119 (2004)
Xu, X. et al. Four-winged dinosaurs from China. Nature 421, 335–340 (2003)
Zhang, F.-C. & Zhou, Z.-H. Leg feathers in an Early Cretaceous bird. Nature 431, 925 (2004)
Xu, X., Zhou, Z.-H. & Prum, R. O. Branched integumental structures in Sinornithosaurus and the origin of feathers. Nature 410, 200–204 (2001)
Zhang, F.-C., Zhou, Z.-H., Xu, X., Wang, X.-L. & Sullivan, C. A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers. Nature 455, 1105–1108 (2008)
Xu, X. & Zhang, F.-C. A new maniraptoran with long metatarsalian feathers from China. Naturwissenschaften 92, 173–177 (2005)
Longrich, N. R. Structure and function of hindlimb feathers in Archaeopteryx lithographica . Paleobiology 32, 417–431 (2006)
Sawyer, R. H. & Knapp, L. W. Avian skin development and the evolutionary origin of feathers. J. Exp. Zool. 298B, 57–72 (2003)
Weidensaul, S. Raptors—The Birds of Prey (Lyons & Burford, 1995)
Xu, X. et al. Origin of flight: could four-winged dinosaurs fly? Nature 438, E3–E4 (2005)
Padian, K. Four-winged dinosaurs, bird precursors, or neither? Bioscience 53, 450–452 (2003)
Jensen, J. A. & Padian, K. Small pterosaurs and dinosaurs from the Uncompahgre Fauna (Brushy Basin Member, Morrison Formation: ?Tithonian), Late Jurassic, western Colorado. J. Paleontol. 63, 364–373 (1989)
Chure, D. J. Koparion douglassi, a new dinosaur from the Morrison Formation (Upper Jurassic) of Dinosaur National Monument: the oldest troodontid (Theropoda: Maniraptora). Brigham Young Univ. Geol. Stud. 40, 11–15 (1994)
Xu, X., Zhao, X.-J. & Clark, J. M. A new therizinosaur from the Lower Jurassic Lufeng Formation of Yunnan, China. J. Vert. Paleontol. 21, 477–483 (2001)
Evans, S. E. & Milner, A. R. in In the Shadow of the Dinosaurs (eds Fraser, N. C. & Sues, H.-D.) 303–321 (Cambridge Univ. Press, 1985)
Acknowledgements
We thank S. Zheng for help with the fieldwork, T. Yu for preparing the specimen, C. Sullivan for editing and commenting on the manuscript, G. Sun, P. Cheng and F. Jin for discussions and comments, and J. Huang for Supplementary Fig. 4i. The fieldwork was supported by grants from the Education Bureau of Liaoning Province (numbers 20060805 and 2008S214) and the Special Fund of Shenyang Normal University. This study was also supported by grants to X.X. from the Chinese Academy of Sciences, the National Natural Science Foundation of China, and Major Basic Research Projects of the Ministry of Science and Technology, China.
Author Contributions D.-Y.H and X.X. designed the project. D.-Y.H., X.X., L.-H.H and L.-J.Z. performed the research. X.X. and D.-Y.H. wrote the manuscript.
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This file contains Supplementary Notes and Data, Supplementary Figures S1-S5 with Legends, Supplementary Character List, Supplementary Matrix and Supplementary References. (PDF 1295 kb)
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Hu, D., Hou, L., Zhang, L. et al. A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus. Nature 461, 640–643 (2009). https://doi.org/10.1038/nature08322
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DOI: https://doi.org/10.1038/nature08322
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