Conulariids and Sphenothallus (Cnidaria, Medusozoa) from the Tonggao Formation (Lower Ordovician, China)


Authors: Van Iten H, Muir LA, Botting JP, Zhang YD, Lin JP

Published in: Bulletin of Geosciences, volume 88, issue 4; pages: 713 - 722; Received 5 December 2012; Accepted in revised form 22 July 2013; Online 14 August 2013

Keywords: Conulariella, Sphenothallus, conulariids, Ordovician, South China,

full text (PDF, 2.17 MB)

Export to RIS



Conulariella sp. and possibly one other conulariid species occur in close association with Sphenothallus sp. in the lower part of the early Floian Tonggao Formation near the town of Sandu, Guizhou Province, China. This is the only known occurrence of Conulariella in Early Ordovician rocks outside of Bohemia (Perunican terrane), and also the first report of Sphenothallus from the Ordovician of China. The apertural margin of Conulariella appears to have been mostly straight, and apertural lappets probably were not present in this genus. Some Tonggao conulariids may have attached to orbiculoid brachiopods in life. Based on the most recent palaeogeographical reconstructions of Gondwana and associated terranes, Conulariella could have had a dispersal path along the Gondwanan margin.


Barnes, C.R. 2004. Ordovician oceans and climate, 72-76. In Webby, B.D., Paris, F., Droser, M.L. & Percival, I.C. (eds) The Great Ordovician Biodiversification Event. Columbia University Press, New York.

Barrande, J. 1867. Systeme silurien du centre de la Boheme. Iere partie, Tome 3. Classe des Mollusques, Ordre des Ptéropodes. 179 pp. Chez l’auteur et editeur, Prague & Paris.

Botting, J.P. & Muir, L.A. 2012. Fauna and ecology of the Holothurian Bed, Llandrindod, Wales, UK (Darriwilian, Middle Ordovician), and the oldest articulated holothurian. Palaeontologia Electronica 15(1), 9A, 1-28,

Bouček, B. 1928. Revision des Conulaires paleozoiques de la Boheme. Palaeontographica Bohemiae 11, 60-108.

Bouček, B. 1939. Conularida, A113-A131. In Schindewolf, O.H. (ed.) Handbuch der Paläozoologie 2A. Gebruder Borntrager, Berlin.

Brabcová, Z. & Kraft, P. 2003. Study of conulariid and related phosphatic conical exoskeletons from the Prague Basin (Czech Republic), 263-266. In Albanesi, G.L., Beresi, M.S. & Peralta, S.H. (eds) Ordovician from the Andes. INSUGEO, Serie Correlación Geológica 17.

Cocks, L.R.M. & Torsvik, T.H. 2013. Earth geography from 500 to 400 million years ago: a faunal and palaeomagnetic review. Journal of the Geological Society, London 159, 631-644.View article

Conway Morris, S. & Chen, M. 1992. Carinachitiids, hexangulaconulariids, and Punctatus: problematical metazoans from the Early Cambrian of South China. Journal of Paleontology 66, 384-406.View article

Fatka, O., Kraft, P. & Szabad, M. 2012. A first report of Sphenothallus Hall, 1847 in the Cambrian of Variscan Europe. Comptes Rendus Palevol 11, 539-547.View article

Fatka, O. & Mergl, M. 2009. The ‘microcontinent’ Perunica: status and story 15 years after conception, 65-101. In Bassett, M.G. (ed.) Early Palaeozoic Peri-Gondwana Terranes: New Insights from Tectonics and Biogeography. Geological Society of London, Special Publication 325.

Gotte, A. 1887. Entwicklungsgeschichte der Aurelia aurita und Cotylorhiza tuberculata. Abhandlungen zur Entwicklungsgeschichte der Tiere. Viertes Heft. 79 pp. Verlag von Leopold Voss, Hamburg & Leipzig.

Hall, J. 1847. Paleontology of New York. Volume 1. Containing descriptions of the organic remains of the Lower Division of the New York system (equivalent to the Lower Silurian rocks of Europe). ix-xxix + 338 pp. C. Van Benthuysen, Albany.

Hatschek, B. 1888. Lehrbuch der Zoologie. Eine morphologische Ubersicht der Thierreiches zur Einfuhrung in das Stadium der Wissenschaft. Erste Liefrung. iv + 304 pp. Fischer, Jena.

Havliček, V., Vaněk, J. & Fatka, O. 1994. Perunica microcontinent in the Ordovician (its position within the Mediterranean Province, series division, benthic and pelagic associations). Sborník geologických věd, Geologie 46, 23-56.

He, Y.X. & Yang, X.H. 1986. Early Cambrian coelenterates from Nanjiang, Sichuan. Bulletin of the Chengdu Institute of Geology and Mineral Resources 7, 31-43. [in Chinese, English summary]

Izawa, M.R.M., Flemming, R.L., Zhan, R. & Jin, J. 2012. Characterization of green clay concretions from the Tonggao Formation, South China: mineralogy, petrogenesis and paleoenvironmental implications. Canadian Journal of Earth Sciences 49, 1018-1026.View article

Kraft, J. & Kraft, P. 1992. The Corymbograptus v-similis Biozone (Klabava Formation, Ordovician of Prague Basin). Folia Musei rerum naturalium Bohemiae occidentalis, Geologica 35, 1-26.

Kraft, J. & Kraft, P. 1993. The Holograptus tardibrachiatus Biozone (Klabava Formation, Ordovician of the Prague Basin). Folia Musei rerum naturalium Bohemiae occidentalis, Geologica 37, 1-34.

Kraft, J. & Kraft, P. 1994. The Azygograptus-Tetragraptus (reclinatus group) Biozone (Klabava Formation, Ordovician of the Prague Basin). Folia Musei rerum naturalium Bohemiae occidentalis, Geologica 40, 1-36.

Lefebvre, B., Ghobadipour, M. & Nardin, E. 2005. Ordovician echinoderms from the Tabas and Damghan regions, Iran: palaeobiogeographical implications. Bulletin de la Société géologique de France 176, 231-242.View article

Leme, J.M., Simoes, M.G., Rodrigues, S.C., Van Iten, H. & Marques, A.C. 2008. Major developments in conulariid research: problems of interpretation and future perspectives. Ameghiniana 45, 407-420.

Li, G.X., Zhu, M.Y., Van Iten, H. & Li, C.W. 2004. Occurrence of the earliest known Sphenothallus Hall in the Lower Cambrian of Southern Shaanxi Province, China. Geobios 37, 229-237.View article

Moore, R.C. & Harrington, H.J. 1956a. Conulata, F54-F66. In Moore, R.C. (ed.) Treatise on Invertebrate Paleontology, Part F, Coelenterata. Geological Society of America & University of Kansas Press, Boulder & Lawrence.

Moore, R.C. & Harrington, H.J. 1956b. Scyphozoa, F27-F38. In Moore, R.C. (ed.) Treatise on Invertebrate Paleontology, Part F, Coelenterata. Geological Society of America & University of Kansas Press, Boulder & Lawrence.

Muir, L.A., Botting, J.P., Van Roy, P., Zhang, Y.D. & Lin, J.P. 2011. A comparison of three early Ordovician Lagerstatten. GSA Abstracts with Programs 43(5), 54.

Muir, L.A., Ng, T.W., Li, X.F., Zhang, Y.D. & Lin, J.P. in press. Palaeoscolecidan worms and a possible nematode from the Early Ordovician of South China. Palaeoworld.

Öpik, A.A. 1927. Beitrage zur Kenntnis der Kukruse-(C2-)Stufe in Eesti. II. Publications of the Geological Institution of the University of Tartu 10, 1-35.

Peterson, K.W. 1979. Development of coloniality in Hydrozoa, 105-139. In Larwood, G. & Rosen, B.R. (eds) Biology and Systematics of Colonial Organisms. Academic Press, New York.

Pillet, J. & Beaulieu, G. 1998. Some Middle and Upper Ordovician conulariids from the Armorican Massif. Bulletin de la Société des Sciences Naturelles de l’Ouest de la France 20, 29-40.

Servais, T. & Sintubin, M. 2009. Avalonia, Armorica, Perunica: terranes, microcontinents, microplates or palaeobiogeographical provinces?, 103-115. In Bassett, M.G. (ed.) Early Palaeozoic Peri-Gondwana terranes: new insights from tectonics and biogeography. Geological Society of London, Special Publications 325.

Torsvik, T.H. 2009. BugPlates: Linking biogeography and palaeogeography. Available online at: (accessed 5 June 2010).

Torsvik, T.H. & Cocks, R.M. 2009. The Lower Paleozoic palaeogeographical evolution of the northeastern and eastern peri-Gondwanan margin from Turkey to New Zealand. Geological Society of London, Special Publications 235, 3-21.View article

Van Iten, H. & Cox, R.S. 1992. Evidence of clonal budding in a radial cluster of Paraconularia crustula (White) (Pennsylvanian: ?Cnidaria). Lethaia 25, 421-426.View article

Van Iten, H., Cox, R.S. & Mapes, R.H. 1992. New data on the morphology of Sphenothallus Hall: implications for its affinities. Lethaia 25, 135-144.View article

Van Iten, H., Fitzke, J.A. & Cox, R.S. 1996. Problematical fossil cnidarians from the Upper Ordovician of the north-central USA. Palaeontology 39, 1037-1064.

Van Iten, H., Konate,M. & Moussa, Y. 2008. Conulariids of the upper Talak Formation (Mississippian, Visean) of northern Niger (West Africa). Journal of Paleontology 82, 192-196.View article

Van Iten, H., Leme, J.M., Simoes, M.G., Marques, A.C. & Collins, A.G. 2006. Reassessment of the phylogenetic affinities of conulariids (?Ediacaran-Triassic) within the subphylum Medusozoa (phylum Cnidaria). Journal of Systematic Palaeontology 4, 109-118.View article

Van Iten, H. & Vyhlasová, Z. 2004. Conulariids, 119-123. In Webby, B.D., Paris, F., Droser, M.L. & Percival, I.C. (eds) The Great Ordovician Biodiversification Event. Columbia University Press, New York.

Van Iten, H., Zhu, M.Y. & Li, G.X. 2010. Redescription of Hexaconularia He and Yang, 1986 (Lower Cambrian, South China): implications for the affinities of conulariid-like small shelly fossils. Palaeontology 53, 191-199.View article

Van Roy, P., Orr, P.J., Botting, J.P., Muir, L.A., Vinther, J., Lefebvre, B., el Hariri, K. & Briggs, D.E.G. 2010. Ordovician faunas of Burgess Shale type. Nature 465, 215-218.View article

Wang, Y., Hao, S.G., Chen, X., Rong, J.Y., Li, G.X., Liu, J. & Xu, H. 2003. Sphenothallus from the Lower Silurian of China. Journal of Paleontology 77, 583-588.View article

Želízko, J.V. 1911. Neue Pteropoden des alteren Palaozoikums Mittelbohmens. Jahrbuch der Kaiserlich-königlichen geologischen Reichsanstalt 61, 41-52.

Želízko, J.V. 1921. Aquivalente der untersilurischen Euloma-Niobe fauna bei Plzenec in Bohmen. Videnskabs-selskabet i Christiania, Matematisk-naturvidenskabelig Klasse, Skrifter 2, 1-27.

Zhan, R. & Jin, J. 2008. Onshore migration of a deep-water brachiopod fauna from the Lower Ordovician Tonggao Formation, Jiangnan Slope, southeastern Guizhou Province, South China. Canadian Journal of Earth Sciences 45, 141-157.View article

Zhu, M.Y., Van Iten, H., Cox, R.S., Zhao, Y.L. & Erdtmann, B.-D. 2000. Occurrence of Byronia Matthew and Sphenothallus Hall in the Lower Cambrian of China. Paläontologische Zeitschrift 74, 227-238.View article