Fossilized gut of the trilobite Lioparia bassleri and the distribution of exceptional preservation in the Cambrian Stage 4-Drumian Manto Formation of North China


Authors: Wang P, Fatka O, Sun Z, Budil P, Gao J

Published in: Bulletin of Geosciences, volume 93, issue 4; pages: 491 - 498; Received 10 October 2017; Accepted in revised form 2 November 2018; Online 29 November 2018

Keywords: trilobites, digestive system, Weifang Biota, soft-tissue preservation, Manto Formation, mid-Cambrian, China,

full text (PDF, 2.26 MB)

Export to RIS



The preservation of digestive structures of trilobites is not unique but recognition and reporting about these remains are rare. Here we describe a specimen of Lioparia bassleri Resser & Endo, 1937, recently collected from the Weifang Biota in the Manto Formation (Cambrian, Miaolingian, Shandong, North China). This exceptionally preserved specimen displays the remains of the digestive system in the cephalon as well as in thoracic and pygidial parts of the body. The morphology of the remains of soft parts preserved in the specimen agrees with a simple digestive tract which included a simple gut tube in the thorax and pygidium associated with four pairs of cephalic gut diverticulae in the glabella. The morphology of the digestive tract of L. bassleri compares favourably with earlier observations on trilobites that had gut diverticulae. The preservation of the digestive system in the studied trilobite confirms that the depositional environment of the Manto Formation was favourable to soft-tissue preservation. Earlier papers on trilobites with preserved soft parts from China are summarized. Exceptional preservation of sponges, palaeoscolecids, radiodonts, bivalved arthropods, trilobites, and echinoderms in the Manto Formation is briefly discussed.


BABCOCK, L.E. 2003. Trilobites in Paleozoic predator-prey systems, and their role in reorganization of early Paleozoic ecosystems, 55-92. In KELLEY, P.A., KOWALEWSKI, M. & HANSEN, T.A. (eds) Predator-prey interactions in the fossil record. Kluwer Academic/Plenum Publishers, New York.View article

BABCOCK, L.E. & PEEL, J.S. 2007. Palaeobiology, taphonomy and stratigraphic significance of the trilobite Buenellus from the Sirius Passet Biota, Cambrian of North Greenland. Memoirs of the Association of Australasian Palaeontologists 34, 401-418.

BLACKWELDER, E. 1907. Stratigraphy of Shantung, 19-58. In WILLIS, B., BLACKWELDER, E. & SARGENT, R.H. (eds) Research in China. Vol. 1, Pt. 1. Descriptive topography and geology. Section 1, northeastern China. Carnegie Institution of Washington, 54(1).

CHEN, J.Y. 2004. The dawn of animal world. 366 pp. Jiangshu Publishing House of Science and Technology, Nanjing. [in Chinese]

CHEN, J.Y. & ZHOU, G.Q. 1997. Biology of the Chengjiang fauna. Bulletin of the National Museum of Natural Science 10, 11-106.

CHEN, J.Y., ZHOU, G.Q., ZHU, M.Y. & YEH, K.Y. 1996. Chengjiang Biota - A unique window of the Cambrian explosion. 222 pp. National Museum of Natural Science, Taichung, Taiwan.

ENGLISH, A.M. & BABCOCK, L.E. 2007. Feeding behaviour of two Ordovician trilobites inferred from trace fossils and non-biomineralised anatomy, Ohio and Kentucky, USA. Memoirs of the Association of Australasian Palaeontologists 34, 537-544.

ERIKSSON, M.E. & TERFELT, F. 2012. Exceptionally preserved Cambrian trilobite digestive System Revealed in 3D by synchrotron-radiation X-ray tomographic microscopy. PLoS ONE 7(4), e35625.View article

FATKA, O., BUDIL, P. & DAVID, M. 2015. Digestive structures in Ordovician trilobites Colpocoryphe and Flexicalymene from the Barrandian area of Czech Republic. Estonian Journal of Earth Sciences 64(4), 255-266.View article

FATKA, O., MERGL, M. & BUDIL, P. 2013b. Preservation of the digestive structures in Harpides (Trilobita) from the Lower Ordovician of the Barrandian area (Czech Republic). Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 270(1), 55-67.View article

FATKA, O., LEROSEY-AUBRIL, R., BUDIL, P. & RAK, Š. 2013a. Fossilised guts from the Upper Ordovician Letná Formation (Prague Basin, Czech Republic). Bulletin of Geosciences 88(1), 95-104.View article

GUO, H., ZAN, S. & LUO, K. 1996. Cambrian stratigraphy and trilobites of eastern Liaoning. 184 pp. Jilin University Press, Changchun. [in Chinese with English abstract]

GUTIÉRREZ-MARCO, J.C., GARCÍA-BELLIDO, D.C., RÁBANO, I. & SÁ, A.A. 2017 Digestive and appendicular soft-parts, with behavioural implications, in a large Ordovician trilobite from the Fezouata Lagerstätte, Morocco. Scientific Reports 7(39728).View article

HOPKINS, M.J., CHEN, F., HU, S. & ZHANG, Z. 2017. The oldest known digestive system consisting of both paired digestive glands and a crop from exceptionally preserved trilobites of the Guanshan Biota (early Cambrian, China). PLoS ONE 12(9), e0184982.View article

HOU, X.G., CLARKSON, E.N.K., YANG, J., ZHANG, X.G., WU, G.Q. & YUAN, Z. 2009. Appendages of early Cambrian Eoredlichia (Trilobita) from the Chengjiang biota, Yunnan, China. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 99(3-4), 213-223.View article

HU, S.X., ZHU, M.Y., LUO, H.L., STEINER, M., ZHAO, F.C., LI, G.X. & ZHANG, Z.F. 2013. The Guangshan Biota. 204 pp. Yunnan Science and Technology Press. Kunming.

HUANG, D.Y. 2012. Eocrinoids from the Cambrian Mantou Formation of Dalian, Liaoning. Chinese Science Bulletin 57(30), 3855-3857.View article

HUANG, D.Y., WANG, Y.Y., GAO, J. & WANG, Y.A. 2012. A new anomalocaridid frontal appendage from the middle Cambrian Mantou Formation of the Tangshan area, Hebei. Acta Palaeontologica Sinica 51(4), 411-415.

LEROSEY-AUBRIL, R., HEGNA, T.A. & OLIVE, S. 2011. Inferring internal anatomy from the trilobite exoskeleton: the relationship between frontal auxiliary impressions and the digestive system. Lethaia 44(2), 166-184.View article

LEROSEY-AUBRIL, R., HEGNA, T.A., KIER, C., BONINO, E., HABERSETZER, J. & CARRÉ, M. 2012a. Exploring the preservation of digestive structures of fossil arthropods: The phosphatisation of trilobite guts in the Weeks Formation Lagerstätte (Cambrian; Utah), 171-172. In ZHAO, Y. L., ZHU, M.Y., PENG, J., GAINES, R.R. & PARSLEY, R.L. (eds) Cryogenian-Ediacaran to Cambrian stratigraphy and paleontology of Guizhou, China. Journal of Guizhou University, Natural Science 29(Sup. 1).

LEROSEY-AUBRIL, R., HEGNA, T.A., KIER, C., BONINO, E., HABERSETZER, J. & CARRÉ, M. 2012b. Controls on gut phosphatisation: the trilobites from the Weeks Formation Lagerstätte (Cambrian; Utah). PLoS ONE 7(3), e32934.View article

LEROSEY-AUBRIL, R., PATERSON, J.R., GIBB, S. & CHATTERTON, B.D.E. 2017. Exceptionally-preserved late Cambrian fossils from the McKay Group (British Columbia, Canada) and the evolution of tagmosis in aglaspidid arthropods. Gondwana Research 42, 264-279.View article

LIN, J.P. 2007. Preservation of the gastrointestinal system in Olenoides (Trilobita) from the Kaili Biota (Cambrian) of Guizhou, China. Memoirs of the Association of Australasian Palaeontologists 33, 179-189.

LIN, J.P., ZHAO, Y.L., RAHMAN, I.A., XIAO, S. & WANG, S. 2010. Bioturbation in Burgess Shale-type Lagerstätten - case study of trace fossil-body fossil association from the Kaili Biota (Cambrian Series 3), Guizhou, China. Palaeogeography, Palaeoclimatology, Palaeoecology 292(1-2), 245-256.View article

LIN, T. 1995. Discovery of late early Cambrian worm from Huainan, Anhui. Acta Palaeontologica Sinica 34(4), 505-508. [in Chinese with English abstract]

LIU, Q., HUANG, D. & GONG, Y. 2012. Sponge fossils from the Cambrian Mantou Formation of Hebi, Henan, Central China. Journal of China University of Geosciences 37(Sup. 2), 129-135. [in Chinese with English abstract]View article

LU, Y. 1957. Trilobites, 249-294. In GU, Z., YANG, Z., XU, J., YIN, Z., YU, J., ZHAO, J., LU, Y. & HOU, Y. (eds) Index fossils of China. Invertebrates, Part 3. Geological Publishing House, Beijing. [in Chinese]

LU, Y. & DONG, N. 1952. Revision of the Cambrian type sections of Shandong. Acta Geologica Sinica 32(3), 1-23. [in Chinese]

MÁNGANO, M.G. 2011. Trace-fossil assemblages in a Burgess Shale-type deposit from the Stephen Formation at Stanley Glacier, Canadian Rocky Mountains: Unravelling ecologic and evolutionary controls, 89-107. In JOHNSTON, P.A. & JOHNSTON, K.J. (eds) Proceedings of the International Conference on the Cambrian Explosion. Palaeontographica Canadiana 31.

MIKULÁŠ, R., FATKA, O. & SZABAD, M. 2012. Paleoecologic implications of ichnofossils associated with slightly skeletonized body fossils, middle Cambrian of the Barrandian area, Czech Republic. Ichnos 19(4), 199-210.View article

PENG, S. 2009. Review on the studies of Cambrian trilobite faunas from Jiangnan Slope Belt, South China, with notes on Cambrian correlation between South and North China. Acta Palaeontologica Sinica 48(3), 437-452. [in Chinese with English abstract]

RESSER, C.E. 1929. New Lower and Middle Cambrian Crustacea. Proceedings of the United States National Museum 76(9), 1-18.View article

RESSER, C.E. & ENDO, R. 1937. Description of the fossils, 103-301. In ENDO, R. & RESSER, C.E. (eds) The Sinian and Cambrian formations and fossils of southern Manchoukuo. Manchurian Science Museum Bulletin 1.

ROBISON, R.A. 1982. Some Middle Cambrian agnostoid trilobites from western north America. Journal of Paleontology 56(1), 132-160.

ROBISON, R.A. 1984. Cambrian Agnostida of North America and Greenland. Part I, Ptychagnostidae. University of Kansas Paleontological Contributions, Paper 109, 1-59.

ROBISON, R.A. & BABCOCK, L.E. 2011. Systematics, paleobiology, and taphonomy of some exceptionally preserved trilobites from Cambrian Lagerstätten of Utah. Kansas University Paleontological Contributions 5, 1-47.

ROBISON, R.A., BABCOCK, L.E. & GUNTHER, V.G. 2015. Exceptional Cambrian fossils from Utah: a window into the Age of Trilobites. Utah Geological Survey, Miscellaneous Publication 15, 1-97.

SHU, D.G., GEYER, G., CHEN, L. & ZHANG, X.L. 1995. Redlichiacean trilobites with preserved soft-parts from the Lower Cambrian Chengjiang fauna (South China). Beringeria, Special Issue 2, 203-241.

SUN, X. 1989. Cambrian agnostids from the North China platform. Palaeontologia Cathayana 4, 53-129.

SUN, Z.X. & YUAN, J.L. 2015. The Weifang Biota from Cambrian Series 3, North China. The 28th Annual Conference of Palaeontological Society of China. Abstract Volume, 40-41. [in Chinese]

SUN, Z.X., WANG, P.L. & YUAN, J.L. 2015. The first complete Tuzoia manchuriensis from the Cambrian Series 3 of Weifang, Shandong Province. Acta Palaeontologica Sinica 54(1), 113-119. [in Chinese with English abstract]

WANG, P.L., SUN, Z.X. & YUAN, J.L. 2014. Preliminary study on an exceptionally well preserved fauna from the Mantou Formation, Cambrian Series 3, Weifang City Shandong, 185-188. In ZHAN, R.B. & HUANG, B. (eds) IGCP 591 Field Workshop 2014, Kunming China, 12-21 August 2014, Extended Summary. Nanjing University Press, Nanjing.

WANG, Y., HUANG, D. & LIEBERMAN, B.S. 2010. New Isoxys (Arthropoda) from the Mantou Formation, Shandong Province. Acta Palaeontologica Sinica 49(3), 398-406. [in Chinese with English abstract]

WANG, Y., LIN, J.P., ZHAO, Y.L. & ORR, P.J. 2009. Palaeoecology of the trace fossil Gordia and its interaction with nonmineralizing taxa from the early Middle Cambrian Kaili Biota, Guizhou Province, South China. Palaeogeography, Palaeoclimatology, Palaeoecology 177(1-2), 141-148.View article

WANG, Y., LU, Y., YANG, J., MU, E. & SHENG, J. 1954. Stratigraphy of Taizi River Valley in Liaoning, 1. Acta Geologica 34(1), 17-64. [in Chinese]

XIANG, L.W., ZHU, Z.L., LI, S.J. & ZHOU, Z.Q. 1999. Chinese stratigraphic lexicon (The Cambrian). 95 pp. Geological Publishing House, Beijing.

YUAN, J., LI, Y., MU, X., LIN, J.P. & ZHU, X. 2012. Trilobite fauna of the Changhia Formation (Cambrian Series 3) from Shandong and adjacent area. Palaeontologica Sinica, New Series B 35, 1-758. [in Chinese with English summary]

YUAN, J.L., ZHAO, Y.L., LI, Y. & HUANG, Y.Z. 2002. Trilobite Fauna of the Kaili Formation (uppermost Lower Cambrian-lower Middle Cambrian) from southeastern Guizhou, South China. 423 pp. Shanghai Science and Technology Press, Shanghai.

ZHANG, X., LIU, W. & ZHAO, I. 2008. Cambrian Burgess Shale-type Lagerstätten in South China: Distribution and significance. Gondwana Research 14(1-2), 255-262.View article

ZHANG, X., HAN, J., ZHANG, Z., LIU, H. & SHU, D. 2003. Reconsideration of the supposed naraoiid larva from the Early Cambrian Chengjiang Lagerstätte, South China. Palaeontology 46(3), 447-465.View article

ZHANG, Z. 1996. Cambrian-Ordovician, 43-44. In ZHANG, Z. & LIU, W. (eds) Lithostratigraphy of Shandong Province. China University of Geosciences Press, Wuhan. [in Chinese]

ZHAO, Y., ZHU, M., BABCOCK, L.E., YUAN, J., PARSLEY, R.L., PENG, J., YANG, X. & WANG, Y. 2005. Kaili Biota: a taphonomic window on diversification of metazoans from the basal Middle Cambrian: Guizhou, China. Acta Geologica Sinica 79(6), 751-765.View article

ZHU, X.J., LEROSEY-AUBRIL, R. & ESTEVE, J. 2014. Gut content fossilization and evidence for detritus feeding habits in an enrolled trilobite from the Cambrian of China. Lethaia 47(1), 66-76.View article