Skip to main content Accessibility help

A new stemmed echinoderm from the Furongian of China and the origin of Glyptocystitida (Blastozoa, Echinodermata)



The Furongian (late Cambrian) is an extremely poorly sampled time in the history of echinoderms, with only few localities yielding complete specimens. Here, we document an exquisitely preserved stemmed echinoderm from the Furongian Sandu Formation in South China that provides important new data illuminating the origin of Glyptocystitida, a common Palaeozoic clade of echinoderms. Sanducystis sinensis n. gen. n. sp. displays an organized theca bearing three circlets of plates (basal, infralateral and lateral), a laterally positioned periproct in the CD interray, a lack of respiratory pectinirhombs and a stem divided in two parts, with expanded inner and outer columnals proximally and narrow, elongate, homeomorphic columnals distally. A phylogenetic analysis places Sanducystis more derived than the columnal-bearing ‘eocrinoid’ Ridersia and sister group of a clade encompassing Macrocystella-like glyptocystitoid rhombiferans from the Furongian onwards. By filling in an important morphological gap, Sanducystis provides a clear understanding of character evolution within Glyptocystitida.


Corresponding author

Author for correspondence:


Hide All
Bather, F. A. 1899. A phylogenetic classification of the Pelmatozoa. British Association for the Advancement of Science 68, 916–23.
Broadhead, T. W. & Sumrall, C. D. 2003. Heterochrony and paedomorphic development of Sprinkleocystis ektopios, new genus and species (Rhombifera, Glyptocystida) from the Middle Ordovician (Carodoc) of Tennessee. Journal of Paleontology 77, 113–20.
Bruguière, J. G. 1791. Tableau Encyclopédique et Méthodique des Trois Règnes de la Nature, contenant l'Helminthologie, ou les Vers Infusoires, les Vers Intestins, les Vers Mollusques, etc., Volume 7. Paris: Panckoucke.
Callaway, C. 1877. On a new area of Upper Cambrian rocks in South Shropshire, with a description of a new fauna. Quarterly Journal of the Geological Society of London 33, 652–72.
Dzik, J. & Orłowski, S. 1993. The Late Cambrian eocrinoid Cambrocrinus . Acta Palaeontologica Polonica 38, 2134.
Han, N. R. & Chen, G. Y. 2008. New stylophorans (Echinodermata) from the Upper Cambrian of Guangxi, South China. Science in China, Series D: Earth Sciences 51, 181–6.
Jell, P. A., Burrett, C. F. & Banks, M. R. 1985. Cambrian and Ordovician echinoderms from eastern Australia. Alcheringa 9, 183208.
Kammer, T. W., Sumrall, C. D., Zamora, S., Ausich, W. I. & Deline, B. 2013. Oral region homologies in Paleozoic crinoids and other plesiomorphic pentaradial echinoderms. PLOS ONE 8 (11), 116.
Kesling, R. V. & Mintz, L. W. 1961. Notes on Lepadocystis moorei (Meek) and Upper Ordovician callocystitid cystoid. University of Michigan Museum of Paleontology Contributions 17, 123–48.
Klein, J. T. 1734. Naturalis dispositio echinodermatum. Accessit lucubratiuncula de aculeis echinorum marinorum, cum spicilegio de belemnitis. Gedani: Schreiber, 79 pp.
Koch, D. L. & Strimple, H. L. 1968. A new Upper Devonian cystoid attached to a discontinuity surface. Iowa Geological Survey Report of Investigations 5, 149.
Lefebvre, B., Sumrall, C. D., Shroat-Lewis, R. A., Reich, M., Webster, G. D., Hunter, A. W., Nardin, E., Rozhnov, S. V., Guensburg, T. E., Touzeau, A., Noailles, F. & Sprinkle, J. 2013. Palaeobiogeography of Ordovician Echinoderms. In Early Palaeozoic Biogeography and Palaeobiogeography (eds Harper, D. A. T. & Servais, T.), pp. 165–90. Geological Society of London, Memoir no. 38.
Lerosey-Aubril, R., Ortega-Hernández, J. & Zhu, X. J. 2013. The first aglaspidid sensu stricto from the Cambrian of China (Sandu Formation, Guangxi). Geological Magazine 150, 565–71.
Paul, C. R. C. 1968. Macrocystella Callaway, the earliest glyptocystitid cystoid. Palaeontology 11, 580600.
Paul, C. R. C. 1988. The phylogeny of the cystoids. In Echinoderm Phylogeny and Evolutionary Biology (eds Paul, C. R. C. & Smith, A. B.), pp. 199213. Oxford: Clarendon Press.
Sprinkle, J. 1973. Morphology and Evolution of Blastozoan Echinoderms. Cambridge (Mass): Harvard University Museum of Comparative Zoology, 283 pp.
Sprinkle, J. & Guensburg, T. E. 2003. Major expansion of echinoderms in the early Late Ordovician (Mohawkian, middle Caradoc) and its possible causes. In Ordovician from the Andes (eds. Albanesi, G. L., Beresi, M. S. & Peralta, S. H.), pp. 327–32. Instituto Superior de Correlación Geológica (INSUGEO), Serie Correlación Geológica 17, 552 pp.
Sumrall, C. D. 2010. A model for elemental homology for the peristome and ambulacra in blastozoan echinoderms. In Echinoderms: Durham (eds Harris, L. G., Böttger, S. A., Walker, C. W. & Lesser, M. P.), pp. 269–76. London: CRC Press.
Sumrall, C. D. & Sprinkle, J. 1999. Early ontogeny of the glyptocystitid rhombiferan Lepadocystis moorei . In Echinoderm Research 1998 (eds Carnevali, M. D. C. & Bonasoro, F.), pp. 409–14. Rotterdam: Balkema.
Sumrall, C. D., Sprinkle, J. & Guensburg, T. E. 1997. Systematics and paleoecology of Late Cambrian echinoderms from the western United States. Journal of Paleontology 71, 1091–109.
Sumrall, C. D. & Waters, J. A. 2012. Universal elemental homology in glyptocystititoids, hemicosmitoids, coronoids and blastoids: steps toward echinoderm phylogenetic reconstruction in derived blastozoan. Journal of Paleontology 86, 956–72.
Sumrall, C. D. & Wray, G. A. 2007. Ontogeny in the fossil record: diversification of body plans and the evolution of “aberrant” symmetry in Paleozoic echinoderms. Paleobiology 33, 149–63.
Swofford, D. L. 2002. PAUP* version 4.0b10 for Macintosh (computer program and documentation). Sunderland, MA: Sinauer Associates.
Ubaghs, G. 1998. Echinodermes nouveaux du Cambrien supérieur de la Montagne Noire. Geobios 31, 809–29.
Zamora, S. 2012. The first Furongian (late Cambrian) echinoderm from the British Isles. Geological Magazine 149, 940–3.
Zamora, S., Lefebvre, B., Álvaro, J. J., Clausen, S., Elicki, O., Fatka, O., Jell, P., Kouchinsky, A., Lin, J-P., Nardin, E., Parsley, R., Rozhnov, S., Sprinkle, J., Sumrall, C. D., Vizcaïno, D. & Smith, A. B. 2013. Cambrian echinoderm diversity and palaeobiogeography. In Early Palaeozoic Biogeography and Palaeobiogeography (eds Harper, D. A. T. & Servais, T.), pp. 157–71. Geological Society of London, Memoir no. 38.
Zamora, S. & Rahman, I. 2014. Deciphering the early evolution of echinoderms with Cambrian fossils. Palaeontology 57 (6), 1105–19.
Zamora, S. & Smith, A. B. 2012. Cambrian stalked echinoderms show unexpected plasticity of arm construction. Proceedings of the Royal Society B 279, 293–8.
Zamora, S., Zhu, X. & Lefebvre, B. 2013. A new Furongian (Cambrian) Echinoderm-Lagerstätte from the Sandu Formation (South China). Cahiers de Biologie Marine 54, 565–9.
Zhan, R.-B., Jin, J., Rong, J.-Y., Zhu, X.-J. & Han, N.-R. 2010. Late Cambrian brachiopods from Jingxi, Guangxi Province, South China. Alcheringa 34, 99133.
Zhou, Z. Y. & Zhen, Y. Y. 2008. Trilobite Record of China. Beijing: Science Press, 402 pp.
Zhou, Z. Y., Zhen, Y. Y., Peng, S. C. & Zhu, X. J. 2008. Notes on Cambrian trilobite biogeography of China. Acta Palaeontologica Sinica 47, 385–92.
Zhu, X. J., Hughes, N. C. & Peng, S. C. 2007. On a new species of Shergoldia Zhang & Jell, 1987 (Trilobita), the family Tsinaniidae and the order Asaphida. Memoirs of the Association of Australasian Palaeontologists 34, 243–53.
Zhu, X. J., Hughes, N. C. & Peng, S. C. 2010. Ventral structure and ontogeny of the late Furongian (Cambrian) trilobite Guangxiaspis guangxiensis Zhou, 1977 and the diphyletic origin of the median suture. Journal of Paleontology 84, 493504.
Zhu, X., Peng, S., Zamora, S., Lefebvre, B. & Chen, G. 2016. Furongian (upper Cambrian) Guole Konservat-Lagerstätte from South China. Acta Geologica Sinica 90, 30–7.
Zhu, X., Zamora, S. & Lefebvre, B. 2014. Morphology and palaeoecology of a new edrioblastoid (Edrioasteroidea) from the Furongian of China. Acta Palaeontologica Polonica 59, 921–6.
Zittel, K. A. 1879. Protozoa, Coelenterata, Echinodermata und Molluscoidea. Handbuch der Paläontologie, Band 1, Paläozoologie. Munich and Leipzig: R. Oldenbourg, 765 pp.



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