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A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus


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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Figure 1: Stratigraphic column of Jurassic and Lowest Cretaceous strata in western Liaoning, showing horizons from which feathered dinosaurs have been described.
Figure 2: Anchiornis huxleyi LPM-B00169.
Figure 3: Selected skeletal elements and associated feathers of LPM-B00169.
Figure 4: Temporally calibrated phylogeny of the Theropoda.

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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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Correspondence to Dongyu Hu or Xing Xu.

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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).

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