A Preliminary Review of Dinosaur Track Assemblages from the Tuchengzi Formation in North China

Lida Xing; Zhang Yu

doi:10.37819/biosis.003.01.0165

A Preliminary Review of Dinosaur Track Assemblages from the Tuchengzi Formation in North China

Lida Xing

Zhang Yu

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Abstract

In northern China, mainly within Western Liaoning, Northern Hebei and Beijing, vertebrate skeletons from the Tuchengzi Formation (Upper Jurassic to Lower Cretaceous) are scarce. However, a large number of new dinosaur track discoveries have made in past 10 years. So far, a total of 19 dinosaur track sites have been found in the Tuchengzi Formation, collectively containing 2,637 individual tracks and representing at least 2,091 trackmakers. This track record has become an important sample for understanding the Tuchengzi dinosaur fauna. The abundant Tuchengzi track record includes six non-avian theropod ichnogenera (Asianopodus, Eubrontes, Grallator, Menglongipus, Therangospodus, and Velociraptorichnus), three bird ichnogenera (Aquatilavipes, Pullornipes, and Koreanaornis), two sauropod ichnogenera (Brontopodus and cf. Parabrontopodus isp.), and a possible ornithopod ichnogenus (cf. Dinehichnu). This assemblage is saurischian-dominated, which is consistent with most Jurassic and Cretaceous track sites in China. The presence of deinonychosaurian tracks is consistent with the rich skeletal record from the Yanliao Biota or Jehol Biota, and the bird tracks are consistent with the record of shorebirds in Jehol Biota. The existence of ornithopod tracks is doubtful and needs further discoveries to be confirmed.

Keywords: Tuchengzi Formation dinosaur tracks ichnofauna Jurassic Cretaceous

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References

Chen, P.J., Li, J.J., Matsukawa, M., Zhang, H., Wang, Q.& Lockley, M.G. (2006). Geological age of dinosaur track-bearing formations in China. Cretaceous Research, 27, 21-32.
Dong, Z.M. (2001). A forefoot of sauropod from the Tuchengzi Formation of Chaoyang area in Liaoning, China. In: Deng T, Wang Y, editors. Proceedings of the 8th annual meeting of the Chinese Society of Vertebrate Paleontology (pp. 29-33). Beijing: China Ocean Press.
Farlow, J.O., Pittman, J.G.& Hawthorne, J.M. (1989). Brontopodus birdi, Lower Cretaceous dinosaur footprints from the U.S. Gulf Coastal Plain. In: Gillette DD, Lockley MG, editors. Dinosaur Tracks and Traces (pp. 371-394). Cambridge: Cambridge University Press.
Fujita, M., Azuma, Y., Lee, Y.N., Lu, J., Dong, Z.M., Noda, Y.& Urano, K. (2007). New theropod track site from the Upper Jurassic Tuchengzi Formation of Liaoning Province, northeastern China. Mem. Fukui Pref. Dinos. Mus, 6, 17-25.
He, J.R., Jiao, R.C., Nan, Y., Hao, C.Y., Zhang, L.L., Ran, S.H., Song, Q.W., Zhang, Y.C., Zhao, D.N.& Cheng, S.Z. (2017). Fossils, paleoclimate and sedimentary environment of Tuchengzi Formation in the Qianjiadian Basin, Yanqing, Beijing -- the first discovery of suspected dinosaur fossils in Yanqing area. Geological Bulletin of China, 36(8),1319-1329.
Ju, S.B., Wang, X.R., Dong, Y., Liu, Y.C., Ji, Y.N.& Ji, Q. (2021). New discovery of dinosaur footprints from Tuchengzi Formation, Xuanhua, Hebei province. Acta Geologica Sinica.
Liu, D., Chiappe, L.M., Zhang, Y.G., Bell, A., Meng, Q.J., Ji, Q.& Wang, X.R. (2014). An advanced, new long-legged bird from the Early Cretaceous of the Jehol Group (northeastern China): Insights into the temporal divergence of modern Birds. Zootaxa, 3884(3), 253-266. doi:10.11646/zootaxa.3884.3.4.
Liu, Y.Q., Kuang, H.W., Peng, N., Xu, H., Chen, J., Xu, J.L., Liu, H.& Zhang, P. (2012). New discovery of dinosaur footprints in the Upper Jurassic–Lower Cretaceous Houcheng Formation at Shangyi, northwestern Hebei Province and its biogeographical implications. J. Palaeogeogr, 14(5), 617-627.
Lockley, M.G, Farlow, J.O.& Meyer, C.A. (1994). Brontopodus and Parabrontopodus ichnogen. nov. and the significance of wide- and narrow-gauge sauropod trackways. Gaia, 10, 135-145.
Lockley, M.G. (1991). Tracking Dinosaurs—A New Look at an Ancient World (pp. 238). New York: Cambridge University Press.
Lockley, M., Matsukawa, M., Ohira, H., Li, J., Wright, J., White, D. & Chen, P. (2006). Bird tracks from Liaoning Province, China: New insights into avian evolution during the Jurassic-Cretaceous transition. Cretaceous Research, 27(1), 33-43.
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
O’Connor, J. K., Gao, K.-Q., & Chiappe, L. M. (2010). A new ornithuromorph (Aves: Ornithothoraces) bird from the Jehol Group indicative of higher-level diversity. Journal of Vertebrate Paleontology, 30(2), 311–321. doi:10.1080/02724631003617498
Pan, Y.H., Sha, J.G., Zhou, Z.H.& Fürsich, F.T. (2013). The Jehol Biota: Definition and distribution of exceptionally preserved relicts of a continental Early Cretaceous ecosystem. Cret Res, 44, 30-38.
Pu, R.G.& Wu, H.Z. (1982). Sporo-Pollen from the Late Mesozoic beds in eastern Heilongjiang province. Bull.shenyang Inst.geol.min.res.chinese Acad.geol, 4, 169-184.
Santos, V.F., Moratalla, J.J.& Royo-Torres, R. (2009). New sauropod trackways from the Middle Jurassic of Portugal. Acta Palaeontol. Pol, 54, 409–422.
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. Vert. Palas, 47, 35-52.
Wan, X.Q., Sun, L.X.& Li, W. (2020). Terrestrial fossil assemblages of Tuchengzi Formation and Jurassic-Cretaceous boundary chronostratigraphy in Yanliao area. Acta Palaeontologica Sinica, 59(1), 1-12.
Wang, S.E., Gao, L.Z., Wan, X.Q.& Song, B. (2013). Ages of Tuchengzi Formation in western Liaoning northern Hebei area in correlation with those of international strata. Geological Bulletin of China, 32(11), 1673-1690 (in Chinese with English abstract).
Wang, W.L., Zhang, H., Zhang, L.J., Zheng, S.L., Yang, F.L., Li, Z.T., Zheng, Y.J.& Ding, Q.H. (2004). Standard sections of Tuchengzi stage and Yixian stage and their stratigraphy, palaeontology and tectonic-volcanic actions (pp. 1-514). Beijing: Geological Publishing House.
Wilson, J.A.& Carrano, M.T. (1999). Titanosaurs and the origin of “wide-gauge” trackways: a biomechanical and systematic perspective on sauropod locomotion. Paleobiology, 25, 252-267.
Xing, L.D.& Lockley, M.G. (2021b). Theropod tracks from the Jurassic–Cretaceous boundary, Tuchengzi Formation, Chengde, China: Review and new observations. Biosis: Biological Systems, 2(2), 271-282.
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., 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., Harris, J.D., Sun, D.H.& Zhao, H.Q. (2009). The earliest known deinonychosaur tracks from the Jurassic–Cretaceous boundary in Hebei, China. Acta Palaeontologica Sinica, 48(4), 662-671.
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.
Xing, L.D., Lockley, M.G., Du, T.M., Zhang, L.J., Klein, H., Romilio, A., Persons, W.S.I.V., 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, 107, 104310.https://doi.org/10.1016/j.cretres.2019.104310
Xing, L.D., Lockley, M.G., Jiang, Z.K., Klein, H., Persons, W.S.I.V., Zhang, P.F.& Wan, X. Q. (2021a). Variation in sauropod trackway pattern from the Tuchengzi Formation (Jurassic-Cretaceous boundary) in western Liaoning, China. Cretaceous Research, 118, 104662.
Xing, L.D., Lockley, M.G., Mao, Z F., Klein, H., Gu, Z.F., Bai, C.D., Qiu, L., Liu, Y.S., Romilio, A., Persons, W.S.I.V.& Wan, X.Q. (2021c). A new dinosaur track site from the earliest Cretaceous (Berriasian) part of the Tuchengzi Formation, Hebei Province, China: Implications for morphology, ontogeny and paleocommunity structure. Palaeogeography, Palaeoclimatology, Palaeoecology, 580, 110619.
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.
Xu, H., Liu, Y.Q., Kuang, H.W.& Peng, N. (2014). Ages of the Tuchengzi Formation in northern China and the terrestrial Jurassic-Cretaceous boundary in China. Earth Sci Front, 21, 203-215.
Xu, H., Liu, Y.Q., Liu, Y.X.& Kuang, H.W. (2011). Stratigraphy, sedimentology and tectonic background of basin evolution of the Late Jurassic-Early Cretaceous Tuchengzi Formation in Yinshan-Yanshan, North China (in Chinese). Earth Sci Front, 18, 88-106
Yabe, H., Inai, Y.& Shikama, T. (1940). Discovery of dinosaurian footprints from the Cretaceous? of Yangshan, Chinchou: preliminary note. Japan: Proceedings of the Imperial Academy, 16, 560-563.
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, J.P., Xing, L.D., Gierliński, G.D., Wu, F.D., Tian, M.Z. & Currie, P. (2012). First record of dinosaur trackways in Beijing, China. Chinese Science Bulletin, 57(2-3), 144-152.
Zhao, X., Cheng, Z.& Xu, X. (1999). The earliest ceratopsian from the Tuchengzi Formation of Liaoning, China. J Vert Paleont, 19, 681-691.
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.
Zhen, S., Li, J., Rao, C., Mateer, N.& Lockley, M.G. (1989). A review of dinosaur footprints in China. In: Gillette, D.D. Lockley, M.G. (Eds.), Dinosaurs: Past and Present (pp. 187-197). Cambridge: Cambridge University Press.
Zheng, S.L., Zhang, W.& Ding, Q.H. (2001). Discovery of fossil plants from Middle-Upper Jurassic Tuchengzi Formation in western Liaoning, China. Acta Palaeontologica Sinica, 40, 67-85.
Zheng, Y.J., Chen, S.W., Ding, Q.H., Li, Y.F., Zhang, J., Wang, J., Li, X.H., Su, F.& Gao, X. Y. (2010). Chronostratigraphy of the Jurassic Tuchengzi Formation in northern Hebei and western Liaoning, China. Geol. Res, 19(4), 292-295.
Zhou, S., O.’Connor, J.K.& Wang, M. (2014). A new species from an ornithuromorph (Aves: Ornithothoraces) dominated locality of the Jehol Biota. Chinese science bulletin, 59(36), 5366-5378. doi: 10.1007/s11434-014-0669-8.
Zhou, Z.H.& Wang, Y. (2010). Vertebrate diversity of the Jehol Biota as compared with other Lagerstätten. Sci China Earth Sci, 53, 1894-1907.

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A Preliminary Review of Dinosaur Track Assemblages from the Tuchengzi Formation in North China. (2022). Biosis: Biological Systems, 3(1), e002. https://doi.org/10.37819/biosis.003.01.0165

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A Preliminary Review of Dinosaur Track Assemblages from the Tuchengzi Formation in North China. (2022). Biosis: Biological Systems, 3(1), e002. https://doi.org/10.37819/biosis.003.01.0165

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[1] Chen, P.J., Li, J.J., Matsukawa, M., Zhang, H., Wang, Q.& Lockley, M.G. (2006). Geological age of dinosaur track-bearing formations in China. Cretaceous Research, 27, 21-32.

[2] Dong, Z.M. (2001). A forefoot of sauropod from the Tuchengzi Formation of Chaoyang area in Liaoning, China. In: Deng T, Wang Y, editors. Proceedings of the 8th annual meeting of the Chinese Society of Vertebrate Paleontology (pp. 29-33). Beijing: China Ocean Press.

[3] Farlow, J.O., Pittman, J.G.& Hawthorne, J.M. (1989). Brontopodus birdi, Lower Cretaceous dinosaur footprints from the U.S. Gulf Coastal Plain. In: Gillette DD, Lockley MG, editors. Dinosaur Tracks and Traces (pp. 371-394). Cambridge: Cambridge University Press.

[4] Fujita, M., Azuma, Y., Lee, Y.N., Lu, J., Dong, Z.M., Noda, Y.& Urano, K. (2007). New theropod track site from the Upper Jurassic Tuchengzi Formation of Liaoning Province, northeastern China. Mem. Fukui Pref. Dinos. Mus, 6, 17-25.

[5] He, J.R., Jiao, R.C., Nan, Y., Hao, C.Y., Zhang, L.L., Ran, S.H., Song, Q.W., Zhang, Y.C., Zhao, D.N.& Cheng, S.Z. (2017). Fossils, paleoclimate and sedimentary environment of Tuchengzi Formation in the Qianjiadian Basin, Yanqing, Beijing -- the first discovery of suspected dinosaur fossils in Yanqing area. Geological Bulletin of China, 36(8),1319-1329.

[6] Ju, S.B., Wang, X.R., Dong, Y., Liu, Y.C., Ji, Y.N.& Ji, Q. (2021). New discovery of dinosaur footprints from Tuchengzi Formation, Xuanhua, Hebei province. Acta Geologica Sinica.

[7] Liu, D., Chiappe, L.M., Zhang, Y.G., Bell, A., Meng, Q.J., Ji, Q.& Wang, X.R. (2014). An advanced, new long-legged bird from the Early Cretaceous of the Jehol Group (northeastern China): Insights into the temporal divergence of modern Birds. Zootaxa, 3884(3), 253-266. doi:10.11646/zootaxa.3884.3.4.

[8] Liu, Y.Q., Kuang, H.W., Peng, N., Xu, H., Chen, J., Xu, J.L., Liu, H.& Zhang, P. (2012). New discovery of dinosaur footprints in the Upper Jurassic–Lower Cretaceous Houcheng Formation at Shangyi, northwestern Hebei Province and its biogeographical implications. J. Palaeogeogr, 14(5), 617-627.

[9] Lockley, M.G, Farlow, J.O.& Meyer, C.A. (1994). Brontopodus and Parabrontopodus ichnogen. nov. and the significance of wide- and narrow-gauge sauropod trackways. Gaia, 10, 135-145.

[10] Lockley, M.G. (1991). Tracking Dinosaurs—A New Look at an Ancient World (pp. 238). New York: Cambridge University Press.

[11] Lockley, M., Matsukawa, M., Ohira, H., Li, J., Wright, J., White, D. & Chen, P. (2006). Bird tracks from Liaoning Province, China: New insights into avian evolution during the Jurassic-Cretaceous transition. Cretaceous Research, 27(1), 33-43.

[12] 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

[13] O’Connor, J. K., Gao, K.-Q., & Chiappe, L. M. (2010). A new ornithuromorph (Aves: Ornithothoraces) bird from the Jehol Group indicative of higher-level diversity. Journal of Vertebrate Paleontology, 30(2), 311–321. doi:10.1080/02724631003617498

[14] Pan, Y.H., Sha, J.G., Zhou, Z.H.& Fürsich, F.T. (2013). The Jehol Biota: Definition and distribution of exceptionally preserved relicts of a continental Early Cretaceous ecosystem. Cret Res, 44, 30-38.

[15] Pu, R.G.& Wu, H.Z. (1982). Sporo-Pollen from the Late Mesozoic beds in eastern Heilongjiang province. Bull.shenyang Inst.geol.min.res.chinese Acad.geol, 4, 169-184.

[16] Santos, V.F., Moratalla, J.J.& Royo-Torres, R. (2009). New sauropod trackways from the Middle Jurassic of Portugal. Acta Palaeontol. Pol, 54, 409–422.

[17] 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. Vert. Palas, 47, 35-52.

[18] Wan, X.Q., Sun, L.X.& Li, W. (2020). Terrestrial fossil assemblages of Tuchengzi Formation and Jurassic-Cretaceous boundary chronostratigraphy in Yanliao area. Acta Palaeontologica Sinica, 59(1), 1-12.

[19] Wang, S.E., Gao, L.Z., Wan, X.Q.& Song, B. (2013). Ages of Tuchengzi Formation in western Liaoning northern Hebei area in correlation with those of international strata. Geological Bulletin of China, 32(11), 1673-1690 (in Chinese with English abstract).

[20] Wang, W.L., Zhang, H., Zhang, L.J., Zheng, S.L., Yang, F.L., Li, Z.T., Zheng, Y.J.& Ding, Q.H. (2004). Standard sections of Tuchengzi stage and Yixian stage and their stratigraphy, palaeontology and tectonic-volcanic actions (pp. 1-514). Beijing: Geological Publishing House.

[21] Wilson, J.A.& Carrano, M.T. (1999). Titanosaurs and the origin of “wide-gauge” trackways: a biomechanical and systematic perspective on sauropod locomotion. Paleobiology, 25, 252-267.

[22] Xing, L.D.& Lockley, M.G. (2021b). Theropod tracks from the Jurassic–Cretaceous boundary, Tuchengzi Formation, Chengde, China: Review and new observations. Biosis: Biological Systems, 2(2), 271-282.

[23] 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.

[24] 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.

[25] Xing, L.D., Harris, J.D., Sun, D.H.& Zhao, H.Q. (2009). The earliest known deinonychosaur tracks from the Jurassic–Cretaceous boundary in Hebei, China. Acta Palaeontologica Sinica, 48(4), 662-671.

[26] 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.

[27] Xing, L.D., Lockley, M.G., Du, T.M., Zhang, L.J., Klein, H., Romilio, A., Persons, W.S.I.V., 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, 107, 104310.https://doi.org/10.1016/j.cretres.2019.104310

[28] Xing, L.D., Lockley, M.G., Jiang, Z.K., Klein, H., Persons, W.S.I.V., Zhang, P.F.& Wan, X. Q. (2021a). Variation in sauropod trackway pattern from the Tuchengzi Formation (Jurassic-Cretaceous boundary) in western Liaoning, China. Cretaceous Research, 118, 104662.

[29] Xing, L.D., Lockley, M.G., Mao, Z F., Klein, H., Gu, Z.F., Bai, C.D., Qiu, L., Liu, Y.S., Romilio, A., Persons, W.S.I.V.& Wan, X.Q. (2021c). A new dinosaur track site from the earliest Cretaceous (Berriasian) part of the Tuchengzi Formation, Hebei Province, China: Implications for morphology, ontogeny and paleocommunity structure. Palaeogeography, Palaeoclimatology, Palaeoecology, 580, 110619.

[30] 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.

[31] Xu, H., Liu, Y.Q., Kuang, H.W.& Peng, N. (2014). Ages of the Tuchengzi Formation in northern China and the terrestrial Jurassic-Cretaceous boundary in China. Earth Sci Front, 21, 203-215.

[32] Xu, H., Liu, Y.Q., Liu, Y.X.& Kuang, H.W. (2011). Stratigraphy, sedimentology and tectonic background of basin evolution of the Late Jurassic-Early Cretaceous Tuchengzi Formation in Yinshan-Yanshan, North China (in Chinese). Earth Sci Front, 18, 88-106

[33] Yabe, H., Inai, Y.& Shikama, T. (1940). Discovery of dinosaurian footprints from the Cretaceous? of Yangshan, Chinchou: preliminary note. Japan: Proceedings of the Imperial Academy, 16, 560-563.

[34] 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.

[35] Zhang, J.P., Xing, L.D., Gierliński, G.D., Wu, F.D., Tian, M.Z. & Currie, P. (2012). First record of dinosaur trackways in Beijing, China. Chinese Science Bulletin, 57(2-3), 144-152.

[36] Zhao, X., Cheng, Z.& Xu, X. (1999). The earliest ceratopsian from the Tuchengzi Formation of Liaoning, China. J Vert Paleont, 19, 681-691.

[37] 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.

[38] Zhen, S., Li, J., Rao, C., Mateer, N.& Lockley, M.G. (1989). A review of dinosaur footprints in China. In: Gillette, D.D. Lockley, M.G. (Eds.), Dinosaurs: Past and Present (pp. 187-197). Cambridge: Cambridge University Press.

[39] Zheng, S.L., Zhang, W.& Ding, Q.H. (2001). Discovery of fossil plants from Middle-Upper Jurassic Tuchengzi Formation in western Liaoning, China. Acta Palaeontologica Sinica, 40, 67-85.

[40] Zheng, Y.J., Chen, S.W., Ding, Q.H., Li, Y.F., Zhang, J., Wang, J., Li, X.H., Su, F.& Gao, X. Y. (2010). Chronostratigraphy of the Jurassic Tuchengzi Formation in northern Hebei and western Liaoning, China. Geol. Res, 19(4), 292-295.

[41] Zhou, S., O.’Connor, J.K.& Wang, M. (2014). A new species from an ornithuromorph (Aves: Ornithothoraces) dominated locality of the Jehol Biota. Chinese science bulletin, 59(36), 5366-5378. doi: 10.1007/s11434-014-0669-8.

[42] Zhou, Z.H.& Wang, Y. (2010). Vertebrate diversity of the Jehol Biota as compared with other Lagerstätten. Sci China Earth Sci, 53, 1894-1907.