The Maotianshan Shales (帽天山页岩) are a series of Early Cambrian sedimentary deposits in the Chiungchussu Formation,[2] famous for their Konservat Lagerstätten, deposits known for the exceptional preservation of fossilized organisms or traces. The Maotianshan Shales form one of some forty Cambrian fossil locations worldwide exhibiting exquisite preservation of rarely preserved, non-mineralized soft tissue, comparable to the fossils of the Burgess Shale of British Columbia, Canada. They take their name from Maotianshan Hill (Chinese: ; pinyin: Màotiānshān; lit. 'Hat Sky Mountain') in Chengjiang County, Yunnan Province, China.

The most famous assemblage of organisms are referred to as the Chengjiang biota for the multiple scattered fossil sites in Chengjiang. The age of the Chengjiang Lagerstätte is locally termed Qiongzhusian, a stage correlated to the late Atdabanian Stage in Siberian sequences of the middle of the Early Cambrian.[3][4] The shales date to ≤518 million years ago.[1] The shales also contain the slightly younger Guanshan biota from Malong District in Yunnan,[3] Kaili biota and Balang fauna in Guizhou, Shipai fauna in Hubei, and sponge faunas of Guizhou and Anhui.[5]

Along with the Burgess Shale, the Maotianshan Shales are remarked as "our best window into the Cambrian 'explosion'",[6] especially on the origin of chordates.[7]

History and scientific significance

Although fossils from the region have been known from the early part of the 10th century, Chengjiang was first recognized for its exquisite states of preservation with the 1984 discovery of the naraoiid Misszhouia, a soft-bodied relative of trilobites. Since then, the locality has been intensively studied by scientists throughout the world, yielding a constant flow of new discoveries and triggering an extensive scientific debate surrounding the interpretation of discoveries. Over this time, taxa have been revised or reassigned to different groups. Interpretations have led to many refinements of the phylogeny of groups[8][9] and even the erection of the new phylum Vetulicolia of primitive deuterostomes.[10][11]

The Chengjiang biota has all the animal groups found in the Burgess Shale; however, since it is ten million years older, it more strongly supports the deduction that metazoans diversified earlier or faster in the early Cambrian than does the Burgess Shale fauna alone. The preservation of an extremely diverse faunal assemblage renders the Maotianshan shale the world's most important for understanding the evolution of early multi-cellular life, particularly the members of phylum Chordata, which includes all vertebrates. The Chengjiang fossils comprise the oldest diverse metazoan assemblage above the Proterozoic-Phanerozoic transition and, thus, the fossil record's best data source for understanding the apparently rapid diversification of life known as the Cambrian Explosion.

One of the most intriguing locations of the Chengjiang biota is the Haiyan Lagerstätte where hundreds of juvenile specimens have been found. This unique location offers insights into the development of most animal groups and as such is a unique deposit in the Cambrian.[12]

IUGS geological heritage site

In respect of 'the Chengjiang fossils represent[ing] an uparalleled record of the fundamentally important rapid diversification of metazoan life in the early Cambrian', the International Union of Geological Sciences (IUGS) included the 'Cambrian Chengjiang fossil site and lagerstätte' in its assemblage of 100 'geological heritage sites' around the world in a listing published in October 2022. The organisation defines an 'IUGS Geological Heritage Site' as 'a key place with geological elements and/or processes of international scientific relevance, used as a reference, and/or with a substantial contribution to the development of geological sciences through history.'[13]

Preservation and taphonomy

Maotianshania cylindrica, a fossil nematomorph worm, Early Cambrian, Chengjiang Maotianshan Shales

The fossils occur in a section of mudstone 50 metres (160 ft) thick in the Yuanshan Member of the Qiongzhusi Formation. The Yuanshan Member is extensive, covering multiple 10,000 square kilometres (3,900 sq mi) of eastern Yunnan Province, where there are many scattered outcrops yielding fossils. Studies of the strata are consistent with a tropical environment with sea level changes and tectonic activity. The region is believed to have been a shallow sea with a muddy bottom. The preserved fauna is primarily benthic and was likely buried by periodic turbidity currents, since most fossils do not show evidence of post-mortem transport. Like the younger Burgess Shale fossils, the paleo-environment enabled preservation of non-mineralized, soft body parts. Fossils are found in thin layers less than an inch thick. The soft parts are preserved as aluminosilicate films, often with high oxidized iron content and often exhibiting exquisite details.

The Chengjiang beds are very deeply weathered, as evidenced by their low specific gravity (i.e., they are very lightweight).[14] Trace fossils are abundant.[15]

Chengjiang fauna

Haikouella lanceolata, Maotianshan Shales, Chengjiang County, Yunnan Province

The Chengjiang biota comprises an extremely diverse faunal assembly, with some 185 species described in the literature as of June 2006. Of these, nearly half are arthropods, few of which had the hard, mineral-reinforced exoskeletons characteristic of all later arthropoda; only about 3% of the organisms known from Chengjiang have hard shells. Most of those are the trilobites (of which there are five species), all of which have been found with traces of legs, antennae, and other soft body parts, an exceedingly rare occurrence in the fossil record. Phylum Porifera (sponges; 15 species) and Priapulida (16 species) are also well represented. Other phyla represented are Brachiopoda, Chaetognatha, Cnidaria, Ctenophora, Echinodermata, Hyolitha, Nematomorpha, Phoronida, and Chordata. Possible molluscs include Wiwaxia and Nectocaris.[16][17]

About one in eight animals are problematic forms of uncertain affinity, some of which may have been evolutionary experiments that survived for only a brief period as benthic environments rapidly changed in the Cambrian. Chengjiang is the richest source of the Lobopodia, a group including many early panarthropods,[18] with six genera represented: Luolishania, Paucipodia, Cardiodictyon, Hallucigenia (also known from the Burgess Shale), Microdictyon, and Onychodictyon.

Perhaps the most important fossils from Chengjiang are eight possible members of phylum Chordata, the phylum to which all vertebrates belong. The most famous is Myllokunmingia, possibly a very primitive agnathid (i.e., jawless fish). Similar to Myllokunmingia is Haikouichthys ercaicunensis, another primitive fish-like animal.

The enigmatic Yunnanozoon lividum is considered to be the earliest hemichordate, possessing many of the characteristic chordate features and providing an anatomical link between invertebrates and chordates. Haikouella lanceolata is described to be the earliest craniate-like chordate. This fish-like animal has many similarities to Y. lividum, but differs in several aspects: It has a discernible heart, dorsal and ventral aorta, gill filaments, and a notochord (neural chord).

At present, there is no agreement as to the systematic placement of the Vetulicola, represented by seven species from Chengjiang. Originally described as crustacean arthropods, the Vetulicola were later erected as a new phylum of primitive deuterostomes by D.G. Shu et al. (Shu 2001). Another researcher places them with the urochordates, based on putative affinity with the phylum Chordata. They are thought to have been swimmers that either were filter feeders or detritivores.

Some two dozen animals from the Chengjiang biota are problematic regarding phylogenetic assignment. Among these, 'Anomalocaris' saron, the alleged predatory terror of the early Cambrian, was the most famous, although that species is later reclassified to Houcaris saron and Innovatiocaris maotianshanensis. Shu (2006) recently described Stromatoveris psygmoglena as a possible bilateran missing link between Ediacaran fronds and Cambrian ctenophores. Cambrocornulitus had a tubicolous shell which probably was biomineralized. It shares some affinities with cornulitids and lophophorates.[19]

The Chengjiang biota is believed to have inhabited a delta front environment rich in oxygen, with high sedimentation rates and major fluctuations in salinity being the main environmental stressors.[20]

Guanshan biota

Located at the Yunnan Province of South China, the Guanshan biota are also Burgess shale-type fossils but slightly younger than the Chengjian biota, and is dated to 515–510 Myr falling within the Cambrian Stage 4.[21][22] Brachiopods are the most abundant species,[23] followed by trilobites. Other species belong to sponges, chancelloriids, cnidarians, ctenophores, priapulids, lobopodians, arthropods, anomalocaridids, hyoliths, molluscs, brachiopods, echinoderms, algae and vetulicolians. There are also the earliest-known eocrinoids, unidentified soft-bodied animals and abundant trace fossils.[5]

The Guanshan biota are regarded as successors of the Chengjian biota,[22] and share many species. The unique species include arthropods like Guangweicaris[24] and Astutuscaris,[25] vetulicolians like Vetulicola gantoucunensis[26] and V. longbaoshanensis.;[27] chordates like Cathaymyrus haikouensis and Zhongxiniscus intermedius.[28]


See also


  1. ^ a b Yang, C.; Li, X.-H.; Zhu, M.; Condon, D. J.; Chen, J. (2018). "Geochronological constraint on the Cambrian Chengjiang biota, South China" (PDF). Journal of the Geological Society. 175 (4): 659–666. Bibcode:2018JGSoc.175..659Y. doi:10.1144/jgs2017-103. ISSN 0016-7649. S2CID 135091168. Archived (PDF) from the original on 2022-10-09. Retrieved 2019-12-12.
  2. ^ Lipps, J. H.; Signor, P. W (1992). Origin and early evolution of the Metazoa. Springer. ISBN 978-0-306-44067-0.
  3. ^ a b Zhang, X.; Liu, W.; Zhao, Y. (2008). "Cambrian Burgess Shale-type Lagerstätten in South China: Distribution and significance". Gondwana Research. 14 (1–2): 255–262. Bibcode:2008GondR..14..255Z. CiteSeerX doi:10.1016/
  4. ^ Rozanov, A. Yu.; Maoyan Zhu, K. L. Pak and P. Yu. Parkhaev (2008). "The 2nd Sino-Russian Symposium on the Lower Cambrian Subdivision". Paleontological Journal. 42 (4): 441–446. Bibcode:2008PalJ...42..441R. doi:10.1134/S0031030108040151. S2CID 129626166.
  5. ^ a b Hu, ShiXue; Zhu, MaoYan; Steiner, Michael; Luo, HuiLin; Zhao, FangChen; Liu, Qi (2010). "Biodiversity and taphonomy of the Early Cambrian Guanshan biota, eastern Yunnan". Science China Earth Sciences. 53 (12): 1765–1773. Bibcode:2010ScChD..53.1765H. doi:10.1007/s11430-010-4086-9. ISSN 1674-7313. S2CID 128882075.
  6. ^ Shu, D. G.; Conway Morris, S.; Han, J.; Chen, L.; Zhang, X. L.; Zhang, Z. F.; Liu, H. Q.; Li, Y.; Liu, J. N. (2001-11-22). "Primitive deuterostomes from the Chengjiang Lagerstätte (Lower Cambrian, China)". Nature. 414 (6862): 419–424. Bibcode:2001Natur.414..419S. doi:10.1038/35106514. ISSN 0028-0836. PMID 11719797. S2CID 4345484.
  7. ^ McMenamin, Mark A. S. (2019). "Cambrian Chordates and Vetulicolians". Geosciences. 9 (8): 354. Bibcode:2019Geosc...9..354M. doi:10.3390/geosciences9080354. ISSN 2076-3263.
  8. ^ Saleh, Farid; Ma, Xiaoya; Guenser, Pauline; Mángano, M. Gabriela; Buatois, Luis A.; Antcliffe, Jonathan B. (2022-08-23). "Probability-based preservational variations within the early Cambrian Chengjiang biota (China)". PeerJ. 10: e13869. doi:10.7717/peerj.13869. ISSN 2167-8359. PMC 9415357. PMID 36032952.
  9. ^ Xianguang, Hou; Ramsköld, Lars; Bergström, Jan (1991). "Composition and preservation of the Chengjiang fauna –a Lower Cambrian soft-bodied biota". Zoologica Scripta. 20 (4): 395–411. doi:10.1111/j.1463-6409.1991.tb00303.x. ISSN 0300-3256.
  10. ^ Shu, Degan (2005). "On the Phylum Vetulicolia". Chinese Science Bulletin. 50 (20): 2342–2354. Bibcode:2005ChSBu..50.2342S. doi:10.1007/BF03183746. ISSN 1001-6538. S2CID 86827605.
  11. ^ Briggs, Derek E. G.; Fortey, Richard A. (2005). "Wonderful strife: systematics, stem groups, and the phylogenetic signal of the Cambrian radiation". Paleobiology. 31 (2_Suppl): 94–112. doi:10.1666/0094-8373(2005)031[0094:WSSSGA]2.0.CO;2. ISSN 0094-8373. S2CID 44066226.
  12. ^ Yang, X.; Kimmig, J.; Zhai, D.; Liu, Y.; Kimmig, S. R.; Peng, S. (2021). "A juvenile-rich palaeocommunity of the lower Cambrian Chengjiang biota sheds light on palaeo-boom or palaeo-bust environments". Nature Ecology & Evolution. 5 (8): 1082–1090. Bibcode:2021NatEE...5.1082Y. doi:10.1038/s41559-021-01490-4. PMID 34183806. S2CID 235674619.
  13. ^ "The First 100 IUGS Geological Heritage Sites" (PDF). IUGS International Commission on Geoheritage. IUGS. Archived (PDF) from the original on 2022-10-27. Retrieved 13 November 2022.
  14. ^ Gaines, R. R.; Briggs, D. E. G.; Yuanlong, Z. (2008). "Cambrian Burgess Shale–type deposits share a common mode of fossilization". Geology. 36 (10): 755–758. Bibcode:2008Geo....36..755G. doi:10.1130/G24961A.1.
  15. ^ Zhang, X. G.; Bergström, J.; Bromley, R. G.; Hou, X. G. (2007). "Diminutive trace fossils in the Chengjiang Lagerstätte". Terra Nova. 19 (6): 407. Bibcode:2007TeNov..19..407Z. doi:10.1111/j.1365-3121.2007.00765.x. S2CID 129833092.
  16. ^ Zhao, F. C.; Smith, M. R.; Yin, Z.-J.; Zeng, H.; Hu, S.-X; Li, G.-X.; Zhu, M.-Y. (2015). "First report of Wiwaxia from the Cambrian Chengjiang Lagerstätte" (PDF). Geological Magazine. 152 (2): 378–382. Bibcode:2015GeoM..152..378Z. doi:10.1017/S0016756814000648. S2CID 129654292.
  17. ^ Smith, M. R. (2013). "Nectocaridid ecology, diversity and affinity: early origin of a cephalopod-like body plan". Paleobiology. 39 (2): 291–321. Bibcode:2013Pbio...39..297S. doi:10.1666/12029. S2CID 85744624.
  18. ^ Smith, M. R.; Ortega Hernández, J. (2014). "Hallucigenia's onychophoran-like claws and the case for Tactopoda". Nature. 514 (7522): 363–366. Bibcode:2014Natur.514..363S. doi:10.1038/nature13576. PMID 25132546. S2CID 205239797.
  19. ^ Xianfeng, Y.; Vinn, O.; Xianguang, H.; Xinglei, T. (2013). "New tubicolous problematic fossil with some "lophophorate" affinities from the Early Cambrian Chengjiang biota in south China". GFF. 135 (2): 184–190. Bibcode:2013GFF...135..184Y. doi:10.1080/11035897.2013.801035. S2CID 129033570. Retrieved 2014-06-11.
  20. ^ Saleh, Farid; Qi, Changshi; Buatois, Luis A.; Mángano, M. Gabriela; Paz, Maximiliano; Vaucher, Romain; Zheng, Quanfeng; Hou, Xian-Guang; Gabbott, Sarah E.; Ma, Xiaoya (23 March 2022). "The Chengjiang Biota inhabited a deltaic environment". Nature Communications. 13 (1): 1569. Bibcode:2022NatCo..13.1569S. doi:10.1038/s41467-022-29246-z. PMC 8943010. PMID 35322027.
  21. ^ Chen, Feiyang; Zhang, Zhifei; Betts, Marissa J.; Zhang, Zhiliang; Liu, Fan (2019). "First report on Guanshan Biota (Cambrian Stage 4) at the stratotype area of Wulongqing Formation in Malong County, Eastern Yunnan, China". Geoscience Frontiers. 10 (4): 1459–1476. Bibcode:2019GeoFr..10.1459C. doi:10.1016/j.gsf.2018.09.010. S2CID 134921385.
  22. ^ a b Zhao, Jun; Li, Yujing; Selden, Paul A.; Cong, Peiyun (2020). "New occurrence of the Guanshan Lagerstätte (Cambrian Series 2, Stage 4) in the Kunming area, Yunnan, southwest China, with records of new taxa". Alcheringa: An Australasian Journal of Palaeontology. 44 (3): 343–355. Bibcode:2020Alch...44..343Z. doi:10.1080/03115518.2020.1781257. ISSN 0311-5518. S2CID 221093834. Archived from the original on 2023-03-18. Retrieved 2023-03-18.
  23. ^ Hu, Shixue; Zhang, Zhifei; Holmer, Lars E.; Skovsted, Christian B. (2010). "Soft-Part Preservation in a Linguliform Brachiopod from the Lower Cambrian Wulongqing Formation (Guanshan Fauna) of Yunnan, South China". Acta Palaeontologica Polonica. 55 (3): 495–505. doi:10.4202/app.2009.1106. ISSN 0567-7920. S2CID 59439966. Archived from the original on 2023-03-18. Retrieved 2023-03-18.
  24. ^ Wu, Yichen; Liu, Jianni (2019). "Anatomy and relationships of the fuxianhuiid euarthropod Guangweicaris from the early Cambrian Guanshan Biota in Kunming, Yunnan, Southwest China revisited". Acta Palaeontologica Polonica. 64. doi:10.4202/app.00542.2018. S2CID 201291723. Archived from the original on 2023-03-18. Retrieved 2023-03-18.
  25. ^ Jiao, De guang; Du, Kunsheng (2022). "A new euarthropod from the Cambrian Stage 4 Guanshan Biota of South China". Acta Palaeontologica Polonica. 67. doi:10.4202/app.00937.2021. S2CID 253068676. Archived from the original on 2023-03-18. Retrieved 2023-03-18.
  26. ^ Huilin, Luo; Xiaoping, Fu; Shixue, Hu; Yong, Li; Liangzhong, Chen; Ting, You; Qi, Liu (2005). "New Vetulicoliids from the Lower Cambrian Guanshan Fauna, Kunming". Acta Geologica Sinica - English Edition. 79 (1): 1–6. Bibcode:2005AcGlS..79....1L. doi:10.1111/j.1755-6724.2005.tb00860.x. S2CID 129031316. Archived from the original on 2023-03-08. Retrieved 2023-03-18.
  27. ^ Li, JinShu; Liu, JianNi; Ou, Qiang (2017). "New observations on Vetulicola longbaoshanensis from the Lower Cambrian Guanshan Biota (Series 2, Stage 4), South China". Science China Earth Sciences. 60 (10): 1795–1804. Bibcode:2017ScChD..60.1795L. doi:10.1007/s11430-017-9088-y. ISSN 1674-7313. S2CID 135037211.
  28. ^ Huilin, Luo; Shixue, Hu; Liangzhong, Chen (2010). "New Early Cambrian Chordates from Haikou, Kunming". Acta Geologica Sinica - English Edition. 75 (4): 345–348. doi:10.1111/j.1755-6724.2001.tb00051.x. S2CID 84505454. Archived from the original on 2022-10-14. Retrieved 2023-03-18.

Further reading

External links

You may also like