Theropod tracks from the Jurassic–Cretaceous boundary, Tuchengzi Formation, Chengde, China: Review and new observations
Keywords:Theropod tracks, Anchisauripus, Grallatorid, Eubrontid, Oviraptosaurs
Previously known theropod dinosaur footprints preserved as natural casts in the Tuchengzi Formation, on a rock wall beside the railway in Nanshuangmiao Village, Shangbancheng Town, Chengde City, were originally assigned to ichnogenus Anchisauripus and tentatively attributed to oviraptosaurs. The assemblage was restudied in more detail by examining the entire assemblage of 55 tracks associated with two horizons. The size range of the 27 measured tracks suggests a more diverse grallatorid–eubrontid assemblage and potentially greater diversity of theropod trackmakers. The label Anchisauripus, which has fallen into disuse in some recent literature, implies trackmakers of medium shape and size in the grallatorid–eubrontid morphological spectrum. However, given the presence of other theropod ichnotaxa in the Jurassic to Early Cretaceous strata of the Tuchengzi Formation and time equivalent units we suggest that explicit reference to the Grallator-Anchisauripus-Eubrontes (GAE) plexus, or simply the term Grallator-Eubrontes plexus be confined to Lower Jurassic assemblages as originally defined and intended. Further study centered on the 16 known Tuchengzi assemblages and older theropod ichnfaunas is necessary to confirm or refute the degree to which grallatorid–eubrontid assemblages from these different epochs are similar or convergent. Even if the tracks are morphologically very similar inferences regarding trackmaker identity are problematic because the same theropodan trackmaker species, genera or even families were not present in both epochs.
Cope, T. D. & Graham, S. A. (2007). Upper crustal response to Mesozoic tectonism in western Liaoning, North China, and implications for lithospheric delamination, Geological Society, London. Special Publications, 280(1), 201-222. https://doi.org/10.1144/SP280.10
Dong, Z.M. (2001). A forefoot of sauropod from the Tuchengzi Formation of Chaoyang area in Liaoning, China. In Deng, T. & Wang, Y. (Eds.), Proceedings of the 8th Annual Meeting of the Chinese Society of Vertebrate Paleontology (pp. 29-33). Beijing: China Ocean Press.
Farlow, J. O., Coroian, D. & Currie, P. J. (2018). Noah’s ravens: interpreting the makers of tridactyl dinosaur Footprints (p.643). Bloomington: Indiana University Press.
Fujita, M., Azuma, Y., Lee, Y.N., Lü, J.C., Dong, Z.M., Noda, Y.& Urano, K. (2007). New theropod track site from the Upper Jurassic Tuchengzi Formation of Liaoning Province, northeastern China. Memoir of the Fukui Prefectural Dinosaur Museum, 6, 17-25.
Gierlinski, G. & Lockley M. G. (2013). A trackmaker for Saurexallopus: ichnological evidence for oviraptosaurid tracks from the Upper Cretaceous of western North America. Chapter 23, In Titus, A. & Loewen, A. (eds). At the Top of the Grand Staircase (p. 526-529). Indiana University Press.
Li, R., Lockley, M. G., Matsukawa & Liu, M. (2015). Important Dinosaur-dominated footprint assemblages from the Lower Cretaceous Tianjialou Formation at the Houzuoshan Dinosaur Park, Junan County, Shandong Province, China. Cretaceous Research 52, 83-100. https://doi.org/10.1016/j.cretres.2014.08.004
Lockley, M.G. (1991). Tracking dinosaurs: a new look at an ancient world. Cambridge, NY: Cambridge University Press.
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.& Xing, L.D. (2015). Flattened fossil footprints: implications for paleobiology. Palaeogeography, Palaeoclimatology, Palaeoecology, 426, 85-94. https://doi.org/10.1016/j.palaeo.2015.03.008
Lockley, M. G., & Hunt, A. P. (1995). Dinosaur Tracks and Other Fossil Footprints of the Western United States (p. 338). Columbia: Columbia University Press.
Lockley, M.G., Hunt, A.P. & Meyer C. (1994). Vertebrate tracks and the ichnofacies concept: implications for paleoecology and palichnostratigraphy. In Donovan S. (Eds.), The paleobiology of trace fossils (p. 241-268). New York (USA): Wiley and Sons.
Lockley, M.G., Meyer, C.A.& Moratalla, J.J. (1998). Therangospodus: trackway evidence for the widespread distribution of a Late Jurassic theropod dinosaur with well-padded feet. Gaia, 15, 339-353.
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., Li, R., Matsukawa, M., Xing, L. Li, J., Liu, M., & Xing, X. (2015). Tracking the yellow dragons: Implications of China's largest dinosaur tracksite (Cretaceous of the Zhucheng area, Shandong Province, China). Palaeogeography, Palaeoclimatology, Palaeoecology, 423, 62-79. https://doi.org/10.1016/j.palaeo.2015.01.028
Lockley, M. G. & Meyer, C. A. (2000). Dinosaur Tracks and other fossil footprints of Europe. Columbia University Press.
Lockley, M. G. & Milner, A. R. C. (In revision). Eubrontes out west. In Farlow, J.O. (ed.) Indiana University Press.
Lockley, M. G., Simmons, E. & Hirschfield, S. (2018). A new Dinosaur track locality in the Late Cretaceous (Maastrichtian) Laramie Formation of Colorado. New Mexico Museum of Natural History and Science, Bulletin, 79, 395-406.
Lockley, M. G. & Xing, L. (In review). A review of the non-avian theropod track record and the implications for the Ontogenetic Niche Shift model. Earth Science Reviews.
Matsukawa, M., Shibata, K., Kukihara, R., Koarai, K. & Lockley, M. G. (2005). Review of Japanese Dinosaur Track localities: implications for ichnotaxonomy, paleogeography and stratigraphic correlation. Ichnos, 12, 201-222. https://doi.org/10.1080/10420940591009231
Matsukawa, M., Lockley, M.G. & Li, J.J. (2006). Cretaceous terrestrial biotas of East Asia, with special reference to dinosaur-dominated ichnofaunas: towards a synthesis. Cretaceous Research 27, 3-21. https://doi.org/10.1016/j.cretres.2005.10.009
Olsen, P.E. (1980). Fossil great lakes of the Newark Supergroup in New Jersey. In Manspeizer, W. (Ed), Field studies in New Jersey geology and guide to field trips (p. 352-398). 52nd Annual Meeting of the New York State Geological Association, Rutgers University, Newark College of Arts and Sciences, Newark.
Olsen, P.E., Smith, J.B.& McDonald, N.G. (1998). Type material of the type species of the classic theropod footprint genera Eubrontes, Anchisauripus and Grallator (Early Jurassic, Hartford and Deerfield basins, Connecticut and Massachusetts, U.S.A.). Journal of Vertebrate Paleontology, 18 (3), 586-601. https://doi.org/10.1080/02724634.1998.10011086
Shikama, T. (1942). Footprints from Chinchou, Manchoukuo, of Jeholosauripus, the Eo-Mesozoic dinosaur. Bulletin of the Central National Museum Manchoukuo, 3, 21-31.
Stoddard, M.C, Yong, E.H., Akkaynak, D., Sheard, C., Tobias, J.A. & Mahadevan, L. (2017). Avian egg shape: Form function and evolution. Science, 356, 1249-1254. https://doi.org/10.1126/science.aaj1945
Sullivan, C., Hone, D.W.E., Cope, T.D., Liu, Y. & Liu, J. (2009). A new occurrence of small theropod tracks in the Tuchengzi Formation of Hebei Province, China. Vertebrata Palasiatica ,47, 35-52.
Xing, L.D., Harris, J.D., Sun, D.H.& Zhao, H.Q. (2009a). The Earliest Known Deinonychosaur Tracks from the Jurassic–Cretaceous boundary in Hebei, China. Acta Palaeontologica Sinica, 48(4), 662-671.
Xing, L.D., Harris, J.D., Feng, X.Y. & Zhang, Z.J. (2009b). Theropod (Dinosauria: Saurischia) tracks from Lower Cretaceous Yixian Formation at Sihetun, Liaoning Province, China and Possible Track Makers. Geological Bulletin of China, 28(6), 705-712.
Xing, L.D., Harris, J.D.& Gierliński, G.D. (2011). Therangospodus and Megalosauripus track assemblage from the Upper Jurassic–Lower Cretaceous Tuchengzi Formation of Chicheng County, Hebei Province, China and Their Paleoecological Implications. Vertebrata PalasiAtica, 49(4), 423-434
Xing, L.D., Gierliński, G.D., Harris, J.D.& Divay, J.D. (2012). A Probable Crouching Theropod Dinosaur Trace from the Jurassic–Cretaceous Boundary in Hebei, China. Geological Bulletin of China, 31(1), 20-25
Xing, L.D., Liu, Y.Q., Kuang, H.W., Klein, H., Zhang, J.P., Burns, M.E., Chen, J., Wang, M.W.& Hu, J. (2014). Theropod and possible ornithopod track assemblages from the Jurassic–Cretaceous boundary Houcheng Formation, Shangyi, northern Hebei, China. Palaeoworld, 23, 200-208. https://doi.org/10.1016/j.palwor.2013.10.002
Xing, L.D., Zhang, J.P., Lockley, M.G., McCrea, R.T., Klein, H., Alcalá, L., Buckley, L.G., Burns, M.E., Kümmell, S.B.& He, Q. (2015). Hints of the early Jehol Biota: important dinosaur footprint assemblages from the Jurassic–Cretaceous Boundary Tuchengzi Formation in Beijing, China. PLOS ONE, 10(4), e0122715. https://doi.org/10.1371/journal.pone.0122715
Xing, L.D., Lockley, M.G.& Zhang, J.P. (2016). Early Cretaceous dinosaur and other tetrapod tracks of southwestern China. Ningbo: Ningbo Publishing House.
Xing, L.D., Lockley, M.G., Kim, K.S., Klein, H., Matsukawa, M., Lim, J.D., Persons, W.S.IV. & Xu, X. (2017). Mid-Cretaceous dinosaur track assemblage from the Tongfosi Formation of China: comparison with the track assemblage of South Korea. Cretaceous Research, 74, 155-164. https://doi.org/10.1016/j.cretres.2017.02.019
Xing, L.D., Klein, H., Qin Z.H., Persons, W.S.IV. & Xu, X. (2018). Lower Cretaceous theropod tracks with the new ichnogenus and combination Lockleypus luanpingensis from the Dabeigou Formation of the Luanping Basin, Hebei Province, China. Palaeoworld, 27, 482-489. https://doi.org/10.1016/j.palwor.2018.09.004
Xing, L.D., Lockley, M.G., Qin, Z.H., Klein, H., Romilio, A., Persons, W.S.IV., Nie, X.& Wan, X.Q. (2019). Dinosaur tracks from the Lower Cretaceous Xiguayuan Formation in the Luanping Basin, Hebei Province, China. Cretaceous Research. https://doi.org/10.1016/j.cretres.2019.06.009
Xing, L.D., Lockley, M.G., Du, T.M., Zhang, L.J., Klein, H., Romilio, A., Persons, W.S.IV., Wang, K., Li, Z.Y.& Wan, X.Q. (2020). Dinosaur tracks from the Jurassic-Cretaceous boundary Tuchengzi Formation (Hebei Province, China) used as building stones in the Chengde imperial summer resort: age, ichnology, and history. Cretaceous Research. https://doi.org/10.1016/j.cretres.2019.104310
Xing, L., Lockley, M.G., Klein, H., Zhang, L., Romilio, A., Persons, W.S.IV, Peng, G., Ye, Y.& Wang, M. (in press). The new ichnotaxon Eubrontes nobitai ichnosp. nov. and other Saurischian tracks from the Lower Cretaceous Jiaguan Formation of Sichuan Province, China
Xing, L., Lockley, M.G., Mao, Z., Gu., Z., Bai, C., Qiu, L., Klein, H., Liu, Y., Wang, M., Romilio, A., Persons, W.S.IV & Wan., X. (in review). A large dinosaur track site from the Jurassic-Cretaceous boundary in the Tuchengzi Formation, Hebei Province, China: morphology, ontogeny and paleocommunity structure.
Xu, H., Liu, Y.Q., Kuang, H.W. & Peng, N. (2014). Age of the Tuchengzi Formation in northern China and the terrestrial Jurassic-Cretaceous boundary in China. Earth Science Frontiers, 21 (2), 203-215 (in Chinese with English abstract).
Xu, X., Zhou, Z.& Wang, X. (2000). The smallest known non-avian theropod dinosaur. Nature, 408, 705-708. https://doi.org/10.1038/35047056
Xu, X., Wang, K., Zhang, K., Ma, Q., Xing, L., Sullivan, C., Hu, D., Cheng, S.& Wang, S. (2012). A gigantic feathered dinosaur from the Lower Cretaceous of China. Nature, 484, 92-95. https://doi.org/10.1038/nature10906
Xu, X. ; Tang, Z.L.& Wang, X.L. (1999). A therizinosauroid dinosaur with integumentary structures from China. Nature, 399 (6734), 350-354. https://doi.org/10.1038/20670
Yabe, H., Inai, Y. & Shikama, T. (1940). Discovery of dinosaurian footprints from the Cretaceous of Yangshan, Chinchou: preliminary note. Proceedings of the Imperial Academy Japan 16, 560-563.
Young, C.C. (1960). Fossil footprints in China. Vertebrata Palasiatica, 4, 53-66. https://doi.org/10.2183/pjab1912.16.560
Zhang, Y.Z., Zhang, J.P., Wu, P., Zhang, X.B. & Bai, S. (2004). Discovery of dinosaur tracks from the Middle-Late Jurassic Tuchengzi Formation in the Chaoyang area, Liaoning Province. International Geology Review, 50, 561-566.
Zhang, H., Wang, X.M., & Liu, X. M. (2008). Constraints on the upper boundary age of the Tiaojishan Formation volcanic rocks in West Liaoning‐North Hebei by LA-ICP-MS dating. Chinese Science Bulletin, 53(22), 3574-3584. https://doi.org/10.1007/s11434-008-0287-4
Zhao, X., Cheng, Z.& Xu, X. (1999). The earliest ceratopsian from the Tuchengzi Formation of Liaoning, China. Journal of Vertebrate Paleontology, 19, 681-691. https://doi.org/10.1080/02724634.1999.10011181
Zhao, X., Cheng, Z., Xu, X.& Makovicky, P.J. (2006). A New Ceratopsian from the Upper Jurassic Houcheng Formation of Hebei, China. Acta Geologica Sinica, 80, 467-473. https://doi.org/10.1111/j.1755-6724.2006.tb00265.x
Zhen, S.N., Li, J.J., Rao, C.G., Mateer, N.J., & Lockley, M.G. (1989). A review of dinosaur footprints in China. In: Gillette, D.D. and Lockley, M.G. (eds) Dinosaur Tracks and Traces (p. 187-197). Cambridge University Press, Cambridge.
Zhou, Z. H. (2014). The Jehol biota, an early cretaceous terrestrial lagerstätte: New discoveries and implications. National Science Review, 1, 543-559. https://doi.org/10.1093/nsr/nwu055