A Review of two Middle Jurassic Theropod Tracksites Discovered in the 1980s from Sichuan Basin


  • Lida Xing China University of Geosciences, Beijing
  • Martin G. Lockley Dinosaur Trackers Research Group, University of Colorado Denver, USA
  • Guangzhao Peng Zigong Dinosaur Museum, China
  • Yong Ye Zigong Dinosaur Museum, China
  • Shan Jiang Zigong Dinosaur Museum, China
  • Anthony Romilio School of Biological Sciences, The University of Queensland, Australia
  • W. Scott Persons IV Mace Brown Museum of Natural History, Department of Geology and Environmental Geosciences, College of Charleston, USA
  • Miaoyan Wang School of the Earth Sciences and Resources, China University of Geosciences, China




Tracks from two sites in the Middle Jurassic Xintiangou Formation in the Wuma Village area, Wuhuang Township, Zizhong have been known and intermittently studied and excavated since the 1980s. The track-bearing surfaces were exposed by a combination of natural weathering and deliberate excavation by residents in a rural agricultural area. The surfaces were used as “threshing floors” for the processing of agricultural crops in an area subject to weathering under a humid sub-tropical climatic regime. Despite the negative effects of weathering on the quality of track preservation, the sites are historically significant in Chinese ichnology as the type areas for many controversially named theropod ichnotaxa. Subsequent researchers challenged the ichnotaxonomy as provincial and over-split, suggesting that many of the tracks, belong to well-known Lower Jurassic ichnogenera. The present study reviews these two sites, providing new information, and confirming that the tracks belong to the ichnogenera Grallator, Eubrontes and Kayentpus which are typical of the globally widespread Lower Jurassic tetrapod biochron. This suggests the Middle Jurassic ichnofauna in Sichuan is like Lower Jurassic ichnofaunas elsewhere. Previous efforts to transfer the ichnospecies to globally, better-known ichnogenera were important in reducing ichnogenus diversity, but did not reduce ichnosepcies diversity. Herein the ichnotaxa are reviewed and it is shown that the ichnospecies names have no utility for comparative study or in assessing assemblage diversity, or biochron composition. It is therefore proposed that the multiple ichnospecies names proposed based on tracks from these two localities can mostly be accommodated under the labels Grallator isp. indet., and Eubrontes isp. indet.


Ahrens, J., Geveci, B., & Law, C. (2005). 36 - ParaView: An End-User Tool for Large-Data Visualization. In: C. D. Hansen, & C. R. Johnson (Eds.), Visualization Handbook. Burlington: Butterworth-Heinemann, 717–731.

Cignoni, P., Callieri, M., Corsini, M., Dellepiane, M., Ganovelli, F., & Ranzuglia, G. (2008). MeshLab: an Open-Source Mesh Processing Tool. In: V. Scarano, R. De Chiara, & U. Erra. Salerno (Eds), Sixth Eurographics Italian Chapter Conference, 129–136.

Hitchcock, E. (1856). On a new fossil fish, and new fossil footmarks. American Journal of Science (series 2), 21, 96–100.

Huang, D. (2019). Jurassic integrative stratigraphy and timescale of China. Science China Earth Sciences, 62, 223–255. https://doi.org/10.1007/s11430-017-9268-7

Li, J. J. (2015). Footprints of Mesozoic Reptilians and Avians-Palaeovertebrata Sinica Fascicle 8 (Serial No. 12), Volume II (Amphibians, Reptilians, and Avians) (in Chinese). Beijing: Science Press, 1–273.

Lockley, M. G. (2009). New perspectives on morphological variation in tridactyl footprints: clues to widespread convergence in developmental dynamics. Geological Quarterly, 53, 415–432.

Lockley, M. G., Matsukawa, M., & Li, J. J. (2003). Crouching theropods in taxonomic jungles: ichnological and ichnotaxonomic investigations of footprints with metatarsal and ischial impressions. Ichnos, 10, 169–177. https://doi.org/10.1080/10420940390256249

Lockley, M. G., & Matsukawa, M. (2009). A review of vertebrate track distributions in East and Southeast Asia. Journal Paleontological Society of Korea, 25, 17–42.

Lockley, M. G., Gierlinski, G. D., & Lucas, S. G. (2011). Kayentapus revisited: notes on the type material and the importance of this theropod footprint ichnogenus. New Mexico Museum of Natural History and Science Bulletin, 53, 330–336.

Lockley, M. G., Li, J. J., Li, R. H., Matsukawa, M., Harris, J. D., & Xing, L. D. (2013). A review of the tetrapod track record in China, with special reference to type ichnospecies: implications for ichnotaxonomy and paleobiology. Acta Geologica Sinica (English edition), 87(1), 1–20. https://doi.org/10.1111/1755-6724.12026

Lockley, M. G., Xing, L. D., Lockwood, J. A. F., & Pond, S. (2014). A review of large Cretaceous ornithopod tracks, with special reference to their ichnotaxonomy. Biological Journal of the Linnean Society, 113, 721–736. https://doi.org/10.1111/bij.12294

Lockley, M. G., Li, J., Xing, L., Guo, B., & Matsukawa, M. (2018). Large theropod and small sauropod trackmakers from the Lower Cretaceous Jingchuan Formation, Inner Mongolia, China. Cretaceous Research, 92, 150–167. https://doi.org/10.1016/j.cretres.2018.07.007

Lucas, S. G. (2007). Tetrapod footprint biostratigraphy and biochronology. Ichnos, 14, 5–38. https://doi.org/10.1080/10420940601006792

Lull, R. S. (1904). Fossil footprints of the Jura-Trias of North America. Memoirs of the Boston Society of Natural History, 5, 461–557.

Milner, A. R. C., Harris, J. D., Lockley, M. G., Kirkland, J. L., & Matthews, N. A. (2009). Bird-like Anatomy, Posture, and Behavior Revealed by an Early Jurassic Theropod Dinosaur Resting Trace. PLoS One, 4(3), e4591. https://doi.org/10.1371/journal.pone.0004591

Xing, L. D., Peng, G. Z., Ye, Y., Lockley, M. G., McCrea, R. T., Currie, P. J., Zhang, J. P., & Burns, M. B. (2014). Large theropod trackway from the Lower Jurassic Zhenzhuchong Formation of Weiyuan County, Sichuan Province, China: Review, new observations and special preservation. Palaeoworld, 23, 285–293. https://doi.org/10.1016/j.palwor.2014.10.010

Xing, L. D., Lockley, M. G., Peng, G. Z., Ye, Y., Zhang, J. P., Matsukawa, M., Klein, H., McCrea, R. T., & Persons, W. S. IV. (2016). Eubrontes and Anomoepus track assemblages from the Middle Jurassic Xiashaximiao Formation of Zizhong County, Sichuan, China: review, ichnotaxonomy and notes on preserved tail traces. New Mexico Museum of Natural History and Science Bulletin, 74, 345–352.

Xing, L. D., Lockley, M. G., Tang, Y. Z., Romilio, A., Xu, T., Li, X. W., Tang, Y., & Li, Y. Z. (2018a). Tetrapod track assemblages from Lower Cretaceous desert facies in the Ordos Basin, Shaanxi Province, China, and their implications for Mesozoic paleoecology. Palaeogeography, Palaeoclimatology, Palaeoecology, 507, 1–14. https://doi.org/10.1016/j.palaeo.2018.05.016

Xing, L. D., Lockley, M. G., Klein, H., Zeng, R., Cai, S. F., Luo, X. C., & Li, C. (2018b). Theropod assemblages and a new ichnotaxon Gigandipus chiappei ichnosp. nov. from the Jiaguan Formation, Lower Cretaceous of Guizhou Province, China. Geoscience Frontiers, 9, 1745–1754. https://doi.org/10.1016/j.gsf.2017.12.012

Xing, L. D., Dai, H., Wei, G. B., Lockley, M. G., Klein, H., Persons, W. S. IV., Wang, M. Y., Jing, S. X., & Hu, H. Q. (2020). The Early Jurassic Kayentapus dominated tracks from Chongqing, China. Historical Biology, https://doi.org/10.1080/08912963.2020.1769093

Yang, X., & Yang, D. (1987). Dinosaur Footprints from Mesozoic of Sichuan Basin. Chengdu City: Science and Technology Publications, 30 (in Chinese).

Yang, C. Y., Jiang, X. K., Li, K., Liu, W. B., & Weni, L. H. (2012). Study of dinosaur footprints (Coelurosauria) in Zizhong, Sichuan, China. Journal of Chengdu University of Technology (Science & Technology Edition), 39(4), 379–387.

Zhen, S. N., Li, J., & Rao, C. (1986). Dinosaur Footprints of Jinning, Yunnan. Memoirs of the Beijing Natural History Museum, 33(5): 1–19.

Zhen, S. N., Li, J., Han, Z., & Yang, X. (1996). The Study of Dinosaur Footprints in China. Chengdu: Sichuan Scientific and Technological Publishing House, 110 (in Chinese with English abstract).




How to Cite

Xing, L., G. Lockley, M. ., Peng , G. ., Ye, Y. ., Jiang , S. ., Romilio , A. ., Persons IV, W. S. ., & Wang , M. . (2021). A Review of two Middle Jurassic Theropod Tracksites Discovered in the 1980s from Sichuan Basin. Biosis: Biological Systems, 2(1), 191-208. https://doi.org/10.37819/biosis.002.01.0094