Paläontologische Zeitschrift

, Volume 84, Issue 2, pp 227–237 | Cite as

The oldest stylophoran echinoderm: a new Ceratocystis from the Middle Cambrian of Germany

  • Imran Alexander RahmanEmail author
  • Samuel Zamora
  • Gerd Geyer
Research Paper


A new stylophoran (Ceratocystis prosthiakida sp. nov.) from the early Middle Cambrian of the Franconian Forest (north-east Bavaria, Germany) is described with the aid of computed tomography, a powerful means of visualizing fossil anatomy. Investigation in this manner enabled previously inaccessible, buried parts of the fossil to be imaged and studied. In addition, a procedure for digitally correcting plate articulations was implemented. Ceratocystis prosthiakida sp. nov. possesses a thin anterior right spine which clearly differentiates it from other species of Ceratocystis. Similar to other Ceratocystis species, C. prosthiakida sp. nov. lived in storm-dominated shallow marine conditions. The species is from the Galgenberg Formation, which has an early Agdzian age in the unified West Gondwanan chronostratigraphy. As a result, it represents the oldest stylophoran known to date.


Ceratocystis Stylophora Echinoderms Cambrian Germany 


Aus dem frühen Mittelkambrium des Frankenwaldes (Nordost-Bayern, Deutschland) wird eine neue Art der Stylophora (Ceratocystis prosthiakida sp. nov.) beschrieben, und zwar mit Hilfe der Computertomographie, einem aussagekräftigen Werkzeug zur Visualisierung anatomischer Merkmale des Fossils. Die Untersuchungen mit dieser Methode erlaubten es, bisher nicht zugängliche, überdeckte Teile des Fossils darzustellen und zu studieren. Zusätzlich wurde eine Prozedur zur digitalen Korrektur von Plattenartikulationen angewandt. Ceratocystis prosthiakida sp. nov. hat einen schlanken anterioren rechten Stachel, der die Art eindeutig von anderen Arten der Gattung Ceratocystis unterscheidet. Ähnlich wie die anderen Arten von Ceratocystis lebte C. prosthiakida sp. nov. in Sturm-dominierten flachmarinen Lebensräumen. Die Art stammt aus der Galgenberg-Formation, die ein Alter des frühen Agdziums nach der chronostratigraphischen Gliederung West-Gondwanas hat. Folglich ist sie die älteste bisher bekannte Stylophore.


Ceratocystis Stylophora Echinodermen Kambrium Deutschland 



We thank Devashish Fuloria and Richard Hamilton (Imperial College London) for XMT analysis and Isabel Pérez (Universidad de Zaragoza) for preparing the photographs used in Fig. 4. We also thank Eladio Liñán (Universidad de Zaragoza) and Mark D. Sutton (Imperial College London) for supervision and comments, and three anonymous reviewers. Imran A. Rahman acknowledges the Natural Environment Research Council (NER/S/A/2005/13593) and Samuel Zamora the Gobierno de Aragón and CAI (European Program) for financial support. This work is a contribution to the Project Consolíder MURERO number CGL2006-12975/BTE from MEC of Spain-FEDER-EU and Grupo Consolidado E-17 from Gobierno de Aragón.

Supplementary material

Video 1

X-ray microtomography (XMT) tomograms of Ceratocystis prosthiakida sp. nov. (MPG 2538 kb)

12542_2009_39_MOESM2_ESM.mpg (3.4 mb)
Video 2 Rotating animation of the original virtual reconstruction of Ceratocystis prosthiakida sp. nov. (MPG 3498 kb)
12542_2009_39_MOESM3_ESM.mpg (4.6 mb)
Video 3 Rotating animation of the corrected virtual reconstruction of Ceratocystis prosthiakida sp. nov. (MPG 4682 kb)


  1. Álvaro, J. Javier, and Daniel Vizcaïno. 1998. Révision biostratigraphique du Cambrien moyen du versant méridional de la Montagne Noire (Languedoc, France). Bulletin de la Société géologique de France 169(2): 233–242.Google Scholar
  2. Álvaro, J. Javier, Bertrand Lefebvre, John H. Shergold, and Daniel Vizcaïno. 2001. The Middle-Upper Cambrian of the southern Montagne Noire. Annales de la Société Géologique du Nord (2ème série) 8(4): 205–211.Google Scholar
  3. Álvaro, J. Javier, Olaf Elicki, Gerd Geyer, Adrian W.A. Rushton, A. Rushton, and John H. Shergold. 2003. Palaeogeographical controls on the Cambrian trilobite immigration and evolutionary patterns reported in the western Gondwana margin. Palaeogeography, Palaeoclimatology, Palaeoecology 195(1–2): 5–35.Google Scholar
  4. Bartels, Christoph, Derek E.G. Briggs, and Günther Brassel. 1998. The Fossils of the Hunsrück Slate. Marine Life in the Devonian. Cambridge: Cambridge University Press.Google Scholar
  5. Bather, Francis A. 1913. Caradocian Cystidea from Girvan. Transactions of the Royal Society of Edinburgh 49(2): 359–529.Google Scholar
  6. Bather, Francis A. 1918. Eocystis, I. Eocystites primaevus. The Geological Magazine 5(2): 49–56.CrossRefGoogle Scholar
  7. Berg-Madsen, Vivianne. 1986. Middle Cambrian cystoid (sensu lato) stem columnals from Bornholm, Denmark. Lethaia 19(1): 67–80.CrossRefGoogle Scholar
  8. Bottjer, David J., Eric H. Davidson, Kevin J. Peterson, and R. Andrew Cameron. 2006. Paleogenomics of echinoderms. Science 314(5801): 956–960.CrossRefGoogle Scholar
  9. Brett, Carlton E., Heather E. Moffat, and Wendy L. Taylor. 1997. Echinoderm taphonomy, taphofacies, and Lagerstätten. Paleontological Society Papers 3: 147–190.Google Scholar
  10. Bruguière, Jean 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., vol. 7. Paris: Panckoucke.Google Scholar
  11. Clausen, Sébastien, and Andrew B. Smith. 2005. Palaeoanatomy and biological affinities of a Cambrian deuterostome (Stylophora). Nature 438(7066): 351–354.CrossRefGoogle Scholar
  12. Courjault-Radé, Pierre, Françoise Debrenne, and Anna Gandin. 1992. Palaeogeographic and geodynamic evolution of the Gondwana continental margins during the Cambrian. Terra Nova 4(6): 657–667.CrossRefGoogle Scholar
  13. Cripps, Anthony P. 1991. A cladistic analysis of the cornutes (stem chordates). Zoological Journal of the Linnean Society 102(4): 333–366.CrossRefGoogle Scholar
  14. David, Bruno, Bertrand Lefebvre, Rich Mooi, and Ronald L. Parsley. 2000. Are homalozoans echinoderms? An answer from the extraxial-axial theory. Paleobiology 26(4): 529–555.CrossRefGoogle Scholar
  15. Dean, William T. 2006. Cambrian stratigraphy and trilobites of the Samur Dağ area, south of Hakkâri, southeastern Turkey. Turkish Journal of Earth Sciences 15(3): 225–257.Google Scholar
  16. Dehm, Richard. 1932. Cystoideen aus dem rheinischen Unterdevon. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, Beilagen-Band, Abteilung A 69: 63–93.Google Scholar
  17. Dehm, Richard. 1934. Untersuchungen an Cystoideen des rheinischen Unterdevons. Sitzungsberichte der mathematisch-naturwissenschaftlichen Abteilung der Bayerischen Akademie der Wissenschaften zu München für 1934: 19–43.Google Scholar
  18. Derstler, Kraig. 1979. Biogeography of the stylophoran carpoids (Echinodermata). In Historical biogeography, plate tectonics and the changing environment, ed. Jane Gray, and ArthurJ Boucot, 91–104. Corvallis: Oregon State University Press.Google Scholar
  19. Domínguez, Patricio, Antone G. Jacobson, and Richard P.S. Jefferies. 2002. Paired gill slits in a fossil with a calcite skeleton. Nature 417(6891): 841–844.CrossRefGoogle Scholar
  20. Elicki, Olaf. 2006. Microbiofacies analysis of Cambrian offshore carbonates from Sardinia (Italy): Environment reconstruction and development of a drowning carbonate platform. Carnets de Géologie CG2006: article 2006/01 (CG2006_A01).
  21. Geyer, Gerd, and Ed Landing. 2001. Middle Cambrian of Avalonian Massachusetts: Stratigraphy and correlation of the Braintree trilobites. Journal of Paleontology 75(1): 116–135.CrossRefGoogle Scholar
  22. Geyer, Gerd, and Ed Landing. 2004. A unified Lower-Middle Cambrian chronostratigraphy for West Gondwana. Acta Geologica Polonica 54(2): 179–218.Google Scholar
  23. Geyer, Gerd, and Heinz Wiefel. 1997. Fränkisch-Thüringisches Schiefergebirge. Courier Forschungsinstitut Senckenberg 200: 56–102.Google Scholar
  24. Geyer, Gerd, Olaf Elicki, Oldřich Fatka, and Anna Żylińska. 2008. Cambrian. In The geology of central Europe, vol. 1, ed. Tom McCann, 155–202. London: Geological Society of London.Google Scholar
  25. Gil Cid, María D., and Patricio Domínguez. 1998. ‘Carpoidea’ and Pelmatozoa from the Middle Cambrian of Zafra (SW Spain). In Echinoderms: San Francisco, ed. Rich Mooi, and Malcolm Telford, 93–98. Rotterdam: Balkema.Google Scholar
  26. Gil Cid, María D., and Patricio Domínguez. 1999. Diversidad de equinodermos y carpoideos en el Cámbrico Inferior y Medio de la Península Ibérica. Coloquios de Paleontología 50: 105–116.Google Scholar
  27. Gill, Edmund D., and Kenneth E. Caster. 1960. Carpoid echinoderms from the Silurian and Devonian of Australia. Bulletins of American Paleontology 41: 1–71.Google Scholar
  28. Gozalo, Rodolfo, Eladio Liñán, Teodoro Palacios, José A. Gámez Vintaned, and Eduardo Mayoral. 2003. The Cambrian of the Iberian Peninsula: An overview. Geologica Acta 1(1): 103–112.Google Scholar
  29. Jaekel, Otto. 1901. Über Carpoideen, eine neue Klasse von Pelmatozoen. Zeitschrift der Deutschen Geologischen Gesellschaft 52: 661–677.Google Scholar
  30. Jefferies, Richard P.S. 1969. Ceratocystis perneri Jaekel—A Middle Cambrian chordate with echinoderm affinities. Palaeontology 12(3): 494–535.Google Scholar
  31. Jefferies, Richard P.S. 1986. The Ancestry of the Vertebrates. British Museum, Natural History: London.Google Scholar
  32. Jefferies, Richard P.S. 2001. The origin and early fossil history of the chordate acustico-lateralis system, with remarks on the reality of the echinoderm–hemichordate clade. In Major events in Early Vertebrate Evolution, ed. Per E. Ahlberg, 40–66. London: Taylor & Francis.Google Scholar
  33. Kim, Dong Hee, Stephen R. Westrop, and Ed Landing. 2002. Middle Cambrian (Acadian Series) conocoryphid and paradoxidid trilobites from the upper Chamberlain’s brook formation, Newfoundland and New Brunswick. Journal of Paleontology 76(5): 822–842.CrossRefGoogle Scholar
  34. Klein, Joseph T. 1734. Naturalis Dispositio Echinodermatum. Accessit Lucubratiuncula de Aculeis Echinorum Marinorum, cum Spicilegio de Belemnitis. Gedani: Schreiber.Google Scholar
  35. Lefebvre, Bertrand. 2000. A new mitrate (Echinodermata, Stylophora) from the Tremadoc of Shropshire (England) and the origin of the Mitrocystitida. Journal of Paleontology 74(5): 890–906.CrossRefGoogle Scholar
  36. Lefebvre, Bertrand. 2003. Functional morphology of stylophoran echinoderms. Palaeontology 46(3): 511–555.CrossRefGoogle Scholar
  37. Lefebvre, Bertrand. 2007. Early palaeozoic palaeobiogeography and palaeoecology of stylophoran echinoderms. Palaeogeography, Palaeoclimatology, Palaeoecology 245(1–2): 156–199.CrossRefGoogle Scholar
  38. Lefebvre, Bertrand, and Oldřich Fatka. 2003. Palaeogeographical and palaeoecological aspects of the Cambro-Ordovician radiation of echinoderms in Gondwanan Africa and peri-Gondwanan Europe. Palaeogeography, Palaeoclimatology, Palaeoecology 195(1–2): 73–97.CrossRefGoogle Scholar
  39. Lefebvre, Bertrand, and Daniel Vizcaïno. 1999. New Ordovician cornutes (Echinodermata, Stylophora) from Montagne Noire, Brittany (France), a revision of the order Cornuta Jaekel 1901. Geobios 32(3): 421–458.CrossRefGoogle Scholar
  40. Lefebvre, Bertrand, Gunther J. Eble, Nicolas Navarro, and Bruno David. 2006. Diversification of atypical Palaeozoic echinoderms: A quantitative survey of patterns of stylophoran disparity, diversity, and geography. Paleobiology 32(3): 483–510.CrossRefGoogle Scholar
  41. Loi, Alfredo, Gian Luigi Pillola, and Francesco Leone. 1995. The Cambrian and early Ordovician of south-western Sardinia. Rendiconti del Seminario della Facoltà di Scienze dell’Università di Cagliari 65: 63–81.Google Scholar
  42. McKerrow, W. Stuart, Christopher R. Scotese, and Martin D. Brasier. 1992. Early Cambrian continental reconstructions. Journal of the Geological Society 149(4): 599–606.CrossRefGoogle Scholar
  43. Mooi, Rich. 2001. Not all written in stone: Interdisciplinary syntheses in echinoderm paleontology. Canadian Journal of Zoology 79(7): 1209–1231.CrossRefGoogle Scholar
  44. Rahman, Imran A., and Sébastien Clausen. 2009. Re-evaluating the palaeobiology and affinities of the Ctenocystoidea (Echinodermata). Journal of Systematic Palaeontology (in press).Google Scholar
  45. Sdzuy, Klaus. 1964. Das Kambrium des Frankenwaldes. 1: Erforschungsgeschichte, Vorkommen und Stratigraphie. Senckenbergiana Lethaea 45(1–4): 201–221.Google Scholar
  46. Sdzuy, Klaus. 1972. Das Kambrium der acadobaltischen Faunenprovinz. Gegenwärtiger Kenntnisstand und Probleme. Zentralblatt für Geologie und Paläontologie. Teil II 1972(1/2): 1–91.Google Scholar
  47. Sdzuy, Klaus. 2000. Das Kambrium des Frankenwaldes. 3. Die Lippertsgrüner Schichten und ihre Fauna. Senckenbergiana Lethaea 79(2): 301–327.CrossRefGoogle Scholar
  48. Sdzuy, Klaus, Eladio Liñán, and Rodolfo Gozalo. 1999. The Leonian Stage (early Middle Cambrian): A unit for Cambrian correlation in the Mediterranean subprovince. Geological Magazine 136(1): 39–48.CrossRefGoogle Scholar
  49. Smith, Andrew B. 2005. The pre-radial history of echinoderms. Geological Journal 40(3): 255–280.CrossRefGoogle Scholar
  50. Sprinkle, James. 1973. Morphology and evolution of blastozoan echinoderms. Harvard University Museum of Comparative Zoology, Special Publication: 1–288.Google Scholar
  51. Sumrall, Colin D., and Gregory A. Wray. 2007. Ontogeny in the fossil record: Diversification of body plans and the evolution of “aberrant” symmetry in Paleozoic echinoderms. Paleobiology 33(1): 149–163.CrossRefGoogle Scholar
  52. Sutton, Mark D., Derek E. G. Briggs, David J. Siveter, and Derek J. Siveter. 2001. Methodologies for the visualization and reconstruction of three-dimensional fossils from the Silurian Herefordshire Lagerstätte. Paleontologica Electronica 4 (1): article 2.
  53. Sutton, Mark D., Derek E.G. Briggs, David J. Siveter, Derek J. Siveter, and Patrick J. Orr. 2002. The arthropod Offacolus kingi (Chelicerata) from the Silurian of Herefordshire, England: Computer based morphological reconstructions and phylogenetic affinities. Proceedings of the Royal Society of London B 269(1497): 1195–1203.CrossRefGoogle Scholar
  54. Ubaghs, Georges. 1967. Le genre Ceratocystis Jaekel (Echinodermata, Stylophora). University of Kansas Paleontological Contributions, Paper 22: 1–16.Google Scholar
  55. Ubaghs, Georges. 1968. Stylophora. In Treatise on invertebrate paleontology. Part S, Echinodemata 1 (2), ed. Raymond C. Moore, S495–S565. Lawrence: The University of Kansas and the Geological Society of America.Google Scholar
  56. Ubaghs, Georges. 1987. Échinodermes nouveaux du Cambrien moyen de la Montagne Noire (France). Annales de Paléontologie 73(1): 1–27.Google Scholar
  57. Ubaghs, Georges, and Richard A. Robison. 1985. A new homoiostelean and a new eocrinoid from the Middle Cambrian of Utah. University of Kansas Paleontological Contributions, Paper 115: 1–24.Google Scholar
  58. Ubaghs, Georges, and Richard A. Robison. 1988. Homalozoan echinoderms of the Wheeler Formation (Middle Cambrian) of Western Utah. University of Kansas Paleontological Contributions, Paper 120: 1–18.Google Scholar
  59. Vizcaïno, Daniel, and Bertrand Lefebvre. 1999. Les échinodermes du Paléozoïque inferieur de Montagne Noire: Biostratigraphie et paléodiversité. Geobios 32(2): 353–364.CrossRefGoogle Scholar
  60. Wurm, Adolf. 1925. Über ein Vorkommen von Mittelcambrium (Paradoxidesschichten) im bayrischen Frankenwald bei Wildenstein südlich Presseck. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, Beilagen-Band, Abteilung B 52: 71–93.Google Scholar
  61. Zamora, Samuel, Rodolfo Gozalo, and Eladio Liñán. 2009. Middle Cambrian gogiid echinoderms from Northeast Spain: Taxonomy, palaeoecology, and palaeogeographic implications. Acta Palaeontologica Polonica 54(2): 253–265.CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2009

Authors and Affiliations

  • Imran Alexander Rahman
    • 1
    Email author
  • Samuel Zamora
    • 2
  • Gerd Geyer
    • 3
    • 4
  1. 1.Department of Earth Science and EngineeringImperial College LondonLondonUK
  2. 2.Área y Museo de Paleontología, Departamento de Ciencias de la TierraUniversidad de ZaragozaZaragozaSpain
  3. 3.Bayerisches Landesamt für Umwelt, Geologischer DienstHofGermany
  4. 4.Institutionen för geovetenskaperUppsala UniversitetUppsalaSweden

Personalised recommendations