Skip to main content Accessibility help

Diversity and species abundance patterns of the Early Cambrian (Series 2, Stage 3) Chengjiang Biota from China

  • Fangchen Zhao (a1), Jean-Bernard Caron (a2), David J. Bottjer (a3), Shixue Hu (a4), Zongjun Yin (a1) and Maoyan Zhu (a1)...


Lagerstätten from the Precambrian–Cambrian transition have traditionally been a relatively untapped resource for understanding the paleoecology of the “Cambrian explosion.” This quantitative paleoecological study is based on 10,238 fossil specimens belonging to 100 animal species, 11 phyla, and 15 ecological categories from the lower Cambrian (Series 2, Stage 3) Chengjiang biota (Mafang locality near Haikou, Yunnan Province, China). Fossils were systematically collected within a 2.5-meter-thick sequence divided into ten stratigraphic intervals. Each interval represents an induced time-averaged assemblage of various event (obrution) beds of unknown duration. Overall, the different fossil assemblages are taxonomically and ecologically similar, suggesting the presence of a single community type recurring throughout the Mafang section. The Mafang community is dominated by epibenthic vagile hunters or scavengers, sessile suspension feeders, and infaunal vagile hunters or scavengers represented primarily by arthropods, brachiopods, and priapulids, respectively. Most species have low abundance and low occurrence frequencies, whereas a few species are numerically abundant and occur frequently. Overall, in structure and ecology the Mafang community is comparable to the Middle Cambrian (Series 3, Stage 5) Burgess Shale biota (Walcott Quarry, Yoho National Park, British Columbia, Canada). This suggests that, despite variations in species identity within taxonomic and ecological groups, the structure and ecology of Cambrian Burgess Shale-type communities remained relatively stable until at least the Middle Cambrian (Series 3, Stage 5) in subtidal to relatively deep-water offshore settings in siliciclastic soft-substrate environments.



Hide All
Alroy, J. 2010a. Fair sampling of taxonomic richness and unbiased estimation of origination and extinction rates. InAlroy, J. and Hunt, G., eds. Quantitative methods in paleobiology. Paleontological Society Papers 16:5580.
Alroy, J. 2010b. The shifting balance of diversity among major marine animal groups. Science 329:11911194.
Bergström, J., Hou, X. G., and Halenius, U. 2007. Gut contents and feeding in the Cambrian arthropod Naraoia. GFF 129:7176.
Briggs, D. E. G., Erwin, D. H., and Collier, F. J. 1994. The fossils of the Burgess Shale. Smithsonian Institution Press, Washington, D.C.
Bush, A. M., and Brame, R. I. 2010. Multiple paleoecological controls on the composition of marine fossil assemblages from the Frasnian (Late Devonian) of Virginia, with a comparison of ordination methods. Paleobiology 36:573591.
Caron, J.-B., and Jackson, D. A. 2006. Taphonomy of the Greater Phyllopod Bed community, Burgess Shale. Palaios 21:451465.
Caron, J.-B., and Jackson, D. A. 2008. Paleoecology of the Greater Phyllopod Bed community, Burgess Shale. Palaeogeography, Palaeoclimatology, Palaeoecology 258:222256.
Caron, J.-B., Conway Morris, S., and Cameron, C. B. 2013. Tubicolous enteropneusts from the Cambrian period. Nature 495:503506.
Chao, A., and Jost, L. 2012. Coverage-based rarefaction and extrapolation: standardizing samples by completeness rather than size. Ecology 93:25332547.
Chen, J. Y. 2004. The dawn of animal world. Jiangsu Science and Technology Press, Nanjing.
Chen, J. Y. 2009. The sudden appearance of diverse animal body plans during the Cambrian explosion. International Journal of Developmental Biology 53:733751.
Chen, J. Y., Zhou, G. Q., Zhu, M. Y., and Yeh, K. Y. 1996. The Chengjiang biota—a unique window on the Cambrian explosion. National Museum of Natural Science, Taichung, Taiwan.
Clausen, S., Hou, X. G., Bergström, J., and Franzén, C. 2010. The absence of echinoderms from the lower Cambrian Chengjiang fauna of China: palaeoecological and palaeogeographical implications. Palaeogeography, Palaeoclimatology, Palaeoecology 294:133141.
Colwell, R. K. 2006. EstimateS: statistical estimation of species richness and shared species from samples, Version 8.0, User's guide and application.
Conway Morris, S. 1986. The community structure of the Middle Cambrian Phyllopod bed (Burgess Shale). Palaeontology 29:423467.
Conway Morris, S. 2008. A redescription of a rare chordate, Metaspriggina walcotti Simonetta and Insom, from the Burgess Shale (Middle Cambrian), British Columbia, Canada. Journal of Paleontology 82:424430.
Conway Morris, S., and Caron, J.-B. 2012. Pikaia gracilens Walcott, a stem-group chordate from the Middle Cambrian of British Columbia. Biological Reviews 87:480512.
Conway Morris, S., and Peel, J. S. 2008. The earliest annelids: Lower Cambrian polychaetes from the Sirius Passet Lagerstätte, Peary Land, North Greenland. Acta Palaeontologica Polonica 53:137148.
Dornbos, S. Q., and Chen, J. Y. 2008. Community palaeoecology of the Early Cambrian Maotianshan Shale biota: ecological dominance of priapulid worms. Palaeogeography, Palaeoclimatology, Palaeoecology 258:200212.
Dunne, J. A., Williams, R. J., Martinez, N. D., Wood, R. A., and Erwin, D. H. 2008. Compilation and network analyses of Cambrian food webs. PLoS Biology 6:06930708.
Erwin, D. H., Laflamme, M., Tweedt, S. M., Sperling, E. A., Pisani, D., and Peterson, K. J. 2011. The Cambrian conundrum: early divergence and later ecological success in the early history of animals. Science 334:10911097.
Gotelli, N. J., and Entsminger, G. L. 2001. EcoSim: null models software for ecology, Version 7.0. Acquired Intelligence Inc. and Kesey-Bear.
Hammer, Ø., Harper, D. A. T., and Ryan, P. D. 2001. PAST: palaeontological statistics package for education and data analysis. Palaeontologica Electronica 4, 1.
Han, J., Shu, D., Zhang, Z., Liu, J., Zhang, X., and Yao, Y. 2006. Preliminary notes on soft-bodied fossil concentrations from the Early Cambrian Chengjiang deposits. Chinese Science Bulletin 51:24822492.
Haug, J., Caron, J.-B., and Haug, C. 2013. Demecology in the Cambrian—synchronized molting in arthropods from the Burgess Shale. BMC Biology 11:64. doi: 10.1186/1741-7007-11-64.
Hentschel, B. T. 1998. Intraspecific variations in δ13C indicate ontogenetic diet changes in deposit-feeding polychaetes. Ecology 79:13571370.
Hou, X. G., Aldridge, R. J., Bergström, J., Siveter, D. J., Siveter, D. J., and Feng, X. H. 2004. The Cambrian fossils of Chengjiang, China: the flowering of early animal life. Blackwell Publishing, Oxford.
Hou, X. G., Siveter, D. J., Aldridge, R. J., and Siveter, D. J. 2008. Collective behavior in an early Cambrian arthropod. Science 322:224.
Hou, X. G., Siveter, D. J., Aldridge, R. J., and Siveter, D. J. 2009. A new arthropod in chain-like associations from the Chengjiang Lagerstätte (Lower Cambrian), Yunnan, China. Palaeontology 52:951961.
Hou, X. G., Aldridge, R. J., Siveter, D. J., Siveter, D. J., Williams, M., Zalasiewicz, J., and Ma, X. Y. 2011. An Early Cambrian hemichordate zooid. Current Biology 21:612616.
Hu, S. X. 2005. Taphonomy and palaeoecology of the Early Cambrian Chengjiang biota from Eastern Yunnan, China. Berliner Paläobiologische Abhandlungen 7:1197.
Hu, S. X., Steiner, M., Zhu, M. Y., Erdtmann, B. -D., Luo, H. L., Chen, L. Z., and Weber, B. 2007. Diverse pelagic predators from the Chengjiang Lagerstätte and the establishment of modern-style pelagic ecosystems in the early Cambrian. Palaeogeography, Palaeoclimatology, Palaeoecology 254:307316.
Hurlbert, S. H. 1971. The nonconcept of species diversity: a critique and alternative parameters. Ecology 52:577589.
Kidwell, S. M. 2013. Time-averaging and fidelity of modern death assemblages: building a taphonomic foundation for conservation palaeobiology. Palaeontology 56:487522.
Luo, H. L., Hu, S. X., Zhang, S. S., and Tao, Y. H. 1997. New occurrence of the Early Cambrian Chengjiang fauna in Haikou, Kunming, Yunnan Province, and study on Trilobitoidea. Acta Geologica Sinica 71:122132.
Magurran, A. E. 2004. Measuring biological diversity. Blackwell Publishing, Oxford.
McAleece, N. 1999. BioDiversity Pro, Version 2.00. The Natural History Museum and the Scottish Association for Marine Science.
Olszewski, T. D. 2004. A unified mathematical framework for the measurement of richness and evenness within and among multiple communities. Oikos 104:277287.
Peters, S. E. 2004. Evenness of Cambrian–Ordovician benthic marine communities in North America. Paleobiology 30:325346.
Shu, D. G. 2008. Cambrian explosion: birth of tree of animals. Gondwana Research 14:219240.
Shu, D. G., Luo, H. L., Conway Morris, S., Zhang, X. L., Hu, S. X., Chen, L., Han, J., Zhu, M., Li, Y., and Chen, L. Z. 1999. Early Cambrian vertebrates from South China. Nature 402:4246.
Shu, D. G., Conway Morris, S., Han, J., Zhang, Z. F., Yasui, K., Janvier, P., Chen, L., Zhang, X. L., Liu, J. N., Li, Y., and Liu, H. Q. 2003. Head and backbone of the Early Cambrian vertebrate Haikouichthys. Nature 421:526529.
Shu, D. G., Conway Morris, S., Han, J., Zhang, Z. F., and Liu, J. N. 2004. Ancestral echinoderms from the Chengjiang deposits of China. Nature 430:422427.
Steiner, M., Zhu, M. Y., Zhao, Y. L., and Erdtmann, B.-D. 2005. Lower Cambrian Burgess Shale-type fossil associations of South China. Palaeogeography, Palaeoclimatology, Palaeoecology 220:129152.
Stirling, G., and Wilsey, B. 2001. Empirical relationships between species richness, evenness, and proportional diversity. American Naturalist 158:286–99.
Vannier, J., and Chen, J. Y. 2002. Digestive system and feeding mode in Cambrian naraoiid arthropods. Lethaia 35:107120.
Zhang, X. G., Hou, X. G., and Bergström, J. 2006. Early Cambrian priapulid worms buried with their lined burrows. Geological Magazine 143:743748.
Zhao, F. C., Caron, J. -B., Hu, S. X., and Zhu, M. Y. 2009. Quantitative analysis of taphofacies and paleocommunities in the Early Cambrian Chengjiang Lagerstätte. Palaios 24:826839.
Zhao, F. C., Zhu, M. Y., and Hu, S. X. 2010. Community structure and composition of the Cambrian Chengjiang biota. Science China (Earth Science) 53:17841799.
Zhao, F. C., Hu, S. X., Caron, J.-B., Zhu, M. Y., Yin, Z. J., and Lu, M. 2012. Spatial variation in the diversity and composition of the Lower Cambrian (Series 2, Stage 3) Chengjiang biota, Southwest China. Palaeogeography. Palaeoclimatology. Palaeoecology 346–347:5465.


Altmetric attention score

Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed