A probable case of heterochrony in the solutan Dendrocystites Barrande, 1887 (Echinodermata: Blastozoa) from the Upper Ordovician of the Prague Basin (Czech Republic) and a revision of the family Dendrocystitidae Bassler, 1938


Authors: Noailles F, Lefebvre B, Kašička L

Published in: Bulletin of Geosciences, volume 89, issue 3; pages: 451 - 476; Received 5 August 2013; Accepted in revised form 15 November 2013; Online 20 March 2014

Keywords: Blastozoa, Dendrocystites, Echinodermata, heterochrony, palaeoecology, Prague Basin, Soluta, Upper Ordovician,

full text (PDF, 3.45 MB)

Export to RIS



The morphology of the Late Ordovician solutan Dendrocystites is reevaluated based on more than 300 specimens from the Letná and Zahořany formations (Prague Basin, Czech Republic). This genus is reported for the first time from the Bohdalec Formation, and its presence is confirmed in the Vinice Formation. The morphology of all specimens of the stratigraphically older species D. barrandei (Sandbian) is identical to that of small to medium-size individuals of D. sedgwicki (Katian). Distinctive characters of D. sedgwicki occur only in the largest specimens, and are all size-related (more asymmetrical thecal outlines, stronger ornamentation, rosetting pattern of thecal plates, proliferation of platelets in the proxistele). Consequently, the transition from D. barrandei to D. sedgwicki is interpreted as the result of heterochronic processes, with the largest individuals of D. sedgwicki displaying hyperadult morphologies (hypermorphosis). Dendrocystites is locally abundant in both the Letná and Zahořany formations, but extremely rare in the deeper deposits of the Vinice and Bohdalec formations. This pattern coincides closely with first order fluctuations of the sea-level in the Prague Basin. The life orientation and implied feeding strategy of Dendrocystites and other solutans are both critically discussed. Several independent lines of evidence suggest that solutans were more likely detritus-feeders. Finally, it is proposed that two morphologically distinct patterns of dististele organization were elaborated independently from the polyplated, undifferentiated stalk-like appendage of Coleicarpus (plesiomorphic condition). Consequently, amajor subdivision of the class Soluta into two main clades (Dendrocystitida ord. nov. and Syringocrinida ord. nov.) is proposed. The monophyly of each order is supported by apomorphies based primarily on the organization of the dististele and the morphology of the periproct. “Dendrocystitesrossicus is reinterpreted as belonging to an unknown genus of syringocrinids, whereas Heckericystis kuckersiana may represent a third species of Dendrocystites.


ALLASINAZ, A. 1999. Invertebrati fossili. 809 pp. Unione Tipografico - Editrice Torinese, Torino.

ARONSON, R.B. & BLAKE, D.B. 1997. Evolutionary paleoecology of dense ophiuroid populations. Paleontological Society Papers 3, 107-119.

ARROYO, F. & LARA, R. 2002. Catálogo de las especies de equinodermos s.l. del Ordovícico espanol. Coloquios de Paleontología 53, 87-108.

AUSICH, W.I., GIL CID, M.D. & DOMINGUEZ ALONSO, P. 2002. Ordovician [Dobrotivian (Llandeilian Stage) to Ashgill] crinoids (phylum Echinodermata) from the Montes de Toledo and Sierra Morena, Spain with implications for paleogeography of Peri-Gondwana. Journal of Paleontology 76, 975-992.View article

BARRANDE, J. 1867. Systeme silurien du centre de la Boheme. 1ere partie: Recherches paléontologiques. Volume 3. Classe des Mollusques. Ordre des Ptéropodes. 179 pp. Bellmann, Prague.

BARRANDE, J. 1872. Systeme silurien du centre de la Boheme. Supplément au Volume 1. Trilobites, Crustacés divers et Poissons. 647 pp. Bellman, Prague.

BARRANDE, J. 1887. Systeme silurien du centre de la Boheme. 1ere partie: Recherches paléontologiques. Volume 7. Classe des Echinodermes. Ordre des Cystidées. 233 pp. Gerhard, Leipzig & Řivnáč, Prague.

BASSLER, R.S. 1915. Bibliographic index of American Ordovician and Silurian fossils. Volume 1. United States National Museum 92, 1-718.View article

BASSLER, R.S. 1938. Pelmatozoa Palaeozoica (Generum et Genotyporum Index et Bibliographia). Fossilium Catalogus I Animalia 83, 1-194.

BASSLER, R.S. & MOODEY, M.W. 1943. Bibliographic and faunal index of Paleozoic pelmatozoan echinoderms. Geological Society of America, Special Papers 45, 1-733.View article

BATHER, F.A. 1899. A phylogenetic classification of the Pelmatozoa. Report of the British Association for the Advancement of Science (Section D), 916-923.View article

BATHER, F.A. 1900. Part 3. The Echinoderma, 1-344. In LANKESTER, E.R. (ed.) A Treatise on Zoology. Adam & Charles Black, London.

BATHER, F.A. 1913. Caradocian Cystidea from Girvan. Transactions of the Royal Society of Edinburgh 49, 359-529. View article

BATHER, F.A. 1926. Cothurnocystis: a study in adaptation. Paläontologische Zeitschrift 7, 1-15.View article

BATHER, F.A. 1928. Dendrocystis in North America. Bulletin of the Geological Survey of Canada 49, 5-8.

BENTON, M.J. 1993. The Fossil Record. 845 pp. Chapman & Hall, London.

BILLINGS, E. 1859. On the Crinoideae of the Lower Silurian Rocks of Canada, 7-66. In GEOLOGICAL SURVEY OF CANADA (ed.) Figures and Descriptions of Canadian Organic Remains. Decade 4. John Lovell, Montreal.

BLAKE, D.B. 2013. Early asterozoan (Echinodermata) diversification: a paleontologic quandary. Journal of Paleontology 87, 353-372.View article

BLAKE, D.B. & GUENSBURG, T.E. 1994. Predation by the Ordovician asteroid Promopalaeaster on a pelecypod. Lethaia 27, 235-239.View article

BRETT, C.E., MOFFAT, H.A. & TAYLOR, W.L. 1997. Echinoderm taphonomy, taphofacies, and Lagerstätten. Paleontological Society Papers 3, 147-190.

BROWER, J.C. 1999. A new pleurocystitid rhombiferan echinoderm from the Middle Ordovician Galena Group of northern Iowa and southern Minnesota. Journal of Paleontology 73, 129-153.

BRUGUIERE, J.G. 1791. Tableau Encyclopédique et Méthodique des Trois Regnes de la Nature, contenant l’Helminthologie, ou les vers Infusoires, les vers Intestins, les vers Mollusques, etc., Volume 7. 180 pp. Panckoucke, Paris.

BRUTHANSOVÁ, J. & KRAFT, P. 2003. Pellets independent of or associated with Bohemian Ordovician body fossils. Acta Palaeontologica Polonica 48, 437-445.

CASTER, K.E. 1967. Homoiostela, 581-627. In MOORE, R.C. (ed.) Treatise on Invertebrate Paleontology. Part S, Echinodermata 1(2). Geological Society of America & University of Texas Press, Boulder & Lawrence.

CASTER, K.E. 1983. A new Silurian carpoid echinoderm from Tasmania and a revision of the Allanicytidiidae. Alcheringa 7, 321-335.View article

CHAUVEL, J. 1941. Recherches sur les cystoides et les carpoides armoricains. Mémoires de la Société géologique et minéralogique de Bretagne 5, 1-286.

CHLUPÁČ, I. 1993. Geology of the Barrandian. A field trip guide. 163 pp. Kramer, Frankfurt-am-Main.

CHLUPÁČ, I., HAVLIČEK, V., KŘÍŽ, J., KUKAL, Z. & ŠTORCH, P. 1998. Palaeozoic of the Barrandian (Cambrian to Devonian). 183 pp. Czech Geological Survey, Prague.

CUÉNOT, L. 1953. Classe des Hétérostélés, 599-606. In PIVETEAU, J. (ed.) Traité de Paléontologie. Masson, Paris.

DALEY, P.E.J. 1992. The anatomy of the solute Girvanicystis batheri (?Chordata) from the Upper Ordovician of Scotland and a new species of Girvanicystis from the Upper Ordovician of South Wales. Zoological Journal of the Linnean Society 105, 353-375.View article

DALEY, P.E.J. 1995. Anatomy, locomotion and ontogeny of the solute Castericystis vali from the Middle Cambrian of Utah. Geobios 28, 585-615.View article

DALEY, P.E.J. 1996. The first solute which is attached as an adult: a Mid-Cambrian fossil from Utah with echinoderm and chordate affinities. Zoological Journal of the Linnean Society 117, 405-440.View article

DAVID, B., LEFEBVRE, B., MOOI, R. & PARSLEY, R.L. 2000. Are homalozoans echinoderms? An answer from the extraxial-axial theory. Paleobiology 26, 529-555.View article

DEHM, R. 1934. Untersuchungen an Cystoideen des Rheinischen Unterdevons. Sitzungsberichte der Bayerischen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Abteilung, Sonderdruck aus dem Jahrgang 1934, 19-34.

DELINE, B. 2008. The first evidence of predatory or parasitic drilling in stylophoran echinoderms. Acta Palaeontologica Polonica 53, 739-743.View article

DOMINGUEZ, P., JEFFERIES, R.P.S. & GIL CID, M.D. 2002. An annotated check-list of genera and species of carpoids. Coloquios de Paleontología 53, 33-68.

DROST, K., GERDES, A., JEFFERIES, T., LINNEMANN, U., STOREY, C. 2011. Provenance of Neoproterozoic and early Paleozoic siliciclastic rocks of the Teplá-Barrandian unit (Bohemian Massif): Evidence from U-Pb detrital zircon ages. Gondwana Research 19, 213-231.View article

DZIK, J. 1999. Evolutionary origin of asymmetry in early metazoan animals, 153-190. In PALYI, G., ZUCCHI, C. & CAGLIOTI, L. (eds) Advances in Biochirality. Elsevier Science, Amsterdam.

EASTON, W.H. 1960. Invertebrate Paleontology. 701 pp. Harper & Brothers, New York.

FATKA, O., LEROSEY-AUBRIL, R., BUDIL, P. & RAK, S. 2013. Fossilised guts in trilobites from the Upper Ordovician Letná Formation (Prague Basin, Czech Republic). Bulletin of Geosciences 88, 95-104.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 Publications 325.

FRIEDRICH, W.P. 1993. Systematik und Funktionsmorphologie mittelkambrischer Cincta (Carpoidea, Echinodermata). Beringeria 7, 1-190.

FUJITA, T. 1992. Dense beds of ophiuroids from the Paleozoic to the Recent: the significance of bathyal populations. Otsuchi Marine Research Centre Reports 18, 25-41.

GIL CID, M.D., DOMINGUEZ ALONSO, P., SILVAN POBES, E. & ESCRIBANO RODENAS, M. 1996. Bohemiaecystis jefferiesi n. sp.; primer Cornuta para el Ordovícico espanol. Estudios Geologicos 52, 313-326.

GILL, E.D. & CASTER, K.E. 1960. Carpoid echinoderms from the Silurian and Devonian of Australia. Bulletins of American Paleontology 41(185), 1-71.

GUENSBURG, T.E. & SPRINKLE, J. 2003. The oldest known crinoids (Early Ordovician, Utah) and a new crinoid plate homology system. Bulletins of American Paleontology 364, 1-43.

GUPTA, V.J. & TERMIER, G. 1978. Lower Palaeozoic Echinodermata from Chandragiri Pass, near Kathmandu, Nepal. Recent Researches in Geology 7, 472-480.

GUTIÉRREZ-MARCO, J.C., RÁBANO, I., SARMIENTO, G.N., ACENOLAZA, G.F., SAN JOSE M.A., PIEREN, A.P., HERRANZ, P., COUTO, H.M. & PIÇARRA, J.M. 1999. Faunal dynamics between Iberia and Bohemia during the Oretanian and Dobrotivian (late Middle - earliest Upper Ordovician), and biogeographic relations with Avalonia and Baltica. Acta Universitatis Carolinae, Geologica 43, 487-490.

HAECKEL, E. 1896. Die Amphorideen und Cystideen. Beiträge zur Morphologie und Phylogenie der Echinodermen. 180 pp. Engelmann, Jena.View article

HAVLÍČEK, V. 1982. Ordovician in Bohemia: development of the Prague Basin and its benthic communities. Sborník geologických věd, Geologie 37, 103-136.

HAVLÍČEK, V. 1989. Climatic changes and development of benthic communities through the Mediterranean Ordovician. Sborník geologických věd, Geologie 44, 79-116.

HAVLÍČEK, V. & FATKA, O. 1992. Ordovician of the Prague Basin (Barrandian area, Czechoslovakia), 461-472. In WEBBY, B.D. & LAURIE, J.R. (eds) Global Perspectives on Ordovician Geology. Balkema, Rotterdam.

HAVLÍČEK, V. & VANĚK, J. 1966. The biostratigraphy of the Ordovician of Bohemia. Sborník geologických věd, Paleontologie 8, 7-69.

HECKER, R.T. 1940. Carpoidea, Eocrinoidea i Ophiocistia nizhnego silura Leningradoskoi oblasti i Estonii. Trudy Paleontologicheskogo instituta Akademii nauk SSSR 9, 5-82.

HOTCHKISS, F.H.C. 2012. Growth zones and extraxial-axial homologies in Asteroidea (Echinodermata). Proceedings of the Biological Society of Washington 125, 106-121.View article

HUNTER, A.W., LEFEBVRE, B., NARDIN, E., RÉGNAULT, S., VAN ROY, P. & ZAMORA, S. 2010. New echinoderm Lagerstätten from the Upper Ordovician of the eastern Anti-Atlas, Morocco, 23-30. In HARRIS, L.G., BÖTTGER, S.A., WALKER, C.W. & LESSER, M.P. (eds) Echinoderms: Durham. Balkema, Leiden.

JAEKEL, O. 1901. Ueber Carpoideen, eine neue Klasse von Pelmatozoen. Zeitschrift der Deutschen geologischen Gesellschaft 52, 661-677.

JAEKEL, O. 1903. Asteriden und Ophiuriden aus dem Silur Böhmens. Zeitschrift der Deutschen geologischen Gesellschaft 55, 106-113.

JAEKEL, O. 1918. Phylogenie und System der Pelmatozoen. Paläontologische Zeitschrift 3, 1-124.View article

JEFFERIES, R.P.S. 1986. The Ancestry of the Vertebrates. 376 pp. British Museum (Natural History), London.

JEFFERIES, R.P.S. 1990. The solute Dendrocystoides scoticus from the Upper Ordovician of Scotland and the ancestry of chordates and echinoderms. Palaeontology 33, 631-679.

JELL, P.A. 1983. Early Devonian echinoderms from Victoria (Rhombifera, Blastoidea and Ophiocistioidea). Memoirs of the Association of Australasian Palaeontologists 1, 209-235.

JELL, P.A. & HOLLOWAY, D. 1982. Anal structure in Rutroclypeus Withers, 1933. Alcheringa 6, 42.View article

KÁCHA, P. & ŠARIČ, R. 2009. Host preferences in Late Ordovician (Sandbian) epibenthic bryozoans: example from the Zahořany Formation of Prague Basin. Bulletin of Geosciences 84, 169-178.View article

KLEIN, J.T. 1734. Naturalis dispositio Echinodermatum. Accessit lucubratiuncula de Aculeis Echinorum Marinorum, cum spicilegio de Belemnitis. 79 pp. Schreiber, Gedani.View article

KOLATA, D.R. 1973. Scalenocystites strimplei, a new Middle Ordovician belemnocystitid solute from Minnesota. Journal of Paleontology 47, 969-974.

KOLATA, D.R., STRIMPLE, H.L. & LEVORSON, C.O. 1977. Revision of the Ordovician carpoid family Iowacystidae. Palaeontology 20, 529-557.

KRÖGER, B., SERVAIS, T. & ZHANG, Y. 2009. The origin and rise of pelagic cephalopods in the Ordovician. Plos One 4, e7262.View article

LEFEBVRE, B. 2000. Homologies in Stylophora: a test of the “calcichordate theory”. Geobios 33, 359-364.View article

LEFEBVRE, B. 2001. Some critical comments on “ankyroids”. Geobios 34, 597-627.View article

LEFEBVRE, B. 2003. Functional morphology of stylophoran echinoderms. Palaeontology 46, 511-555.View article

LEFEBVRE, B. 2007. Early Palaeozoic palaeobiogeography and palaeoecology of stylophoran echinoderms. Palaeogeography, Palaeoclimatology, Palaeoecology 245, 156-199.View article

LEFEBVRE, B., DERSTLER, K. & SUMRALL, C.D. 2012. A reinterpretation of the solutan Plasiacystis mobilis (Echinodermata) from the Middle Ordovician of Bohemia, 287-306. In KROH, A. & REICH, M. (eds) Echinoderm Research 2010: Proceedings of the Seventh European Conference on Echinoderms, Göttingen, Germany, 2-9 October 2010. Zoosymposia 7.

LEFEBVRE, B. & FATKA, O. 2003. Palaeogeographical and palaeoecological aspects of the Cambro-Ordovician radiation of echinoderms in Gondwanan Africa and peri-Gondwanan Europe. Palaeogeography, Palaeoclimatology, Palaeoecology 195, 73-97.View article

LEFEBVRE, B., NARDIN, E., HUNTER, A.W. & RÉGNAULT, S. 2007. Les échinodermes de l’Ordovicien supérieur de l’Anti-Atlas (Maroc) : biostratigraphie et paléodiversité. Africa Geoscience Review 14, 123-148.

LEFEBVRE, B., NOAILLES, F., FRANZIN, B., RÉGNAULT, S., NARDIN, E., HUNTER, A.W., ZAMORA, S., VAN ROY, P., EL HARIRI, K. & LAZREQ, N. 2010. Les gisements a échinodermes de l’Ordovicien supérieur de l’Anti-Atlas oriental (Maroc): un patrimoine scientifique exceptionnel a préserver. Bulletin de l’Institut Scientifique, Rabat 32, 1-17.

LEFEBVRE, B., ROZHNOV, S.V. & FATKA, O. 2005. Lower Palaeozoic “carpoid” echinoderms of Baltica: palaeoecological and palaeobiogeographic implications, 68-69. In KOREN, T., EVDOKIMOVA, I. & TOLMACHEVA, T. (eds) Abstracts, The Sixth Baltic Stratigraphical Conference, August 23-25, St. Petersburg.

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. in press. Palaeobiogeography of Ordovician echinoderms. In HARPER, D.A.T. & SERVAIS, T. (eds) Early Palaeozoic Palaeobiogeography and Palaeogeography. Geological Society of London, Memoir 38.

MARTÍ MUS, M. 2009. Primeros Equinodermos, 450-457. In MARTÍNEZ-CHACÓN, M.L. & RIVAS, P. (eds) Paleontologia de Invertebrados. Universidad de Oviedo, Oviedo.

MORET, L. 1958. Manuel de Paléontologie Animale. 771 pp. Masson, Paris.

MIKULÁŠ, R. 1999. Joint occurrences of body- and trace-fossil communities (Ordovician, Barrandian area, Czech Republic). Journal of the Czech Geological Society 44, 69-78.

MIKULÁŠ, R., PETR, V. & PROKOP, R. 1995. The first occurrence of a “brittlestar bed” (Echinodermata, Ophiuroidea) in Bohemia (Ordovician, Czech Republic). Věstník Českého geologického ústavu 70, 17-24.

MILLER, S.A. & GURLEY, W.F.E. 1894. New genera and species of Echinodermata. Illinois State Museum of Natural History 5, 5-53.

MOOI, R., DAVID, B. & MARCHAND, D. 1994. Echinoderm skeletal homologies: classical morphology meets modern phylogenetics, 87-95. In DAVID, B., GUILLE, A., FÉRAL, J.P. & ROUX, M. (eds) Echinoderms through Time. Balkema, Rotterdam.

NARDIN, E., LEFEBVRE, B., DAVID, B. & MOOI, R. 2009. La radiation des échinodermes au Paléozoique inférieur, l’exemple des blastozoaires. Comptes Rendus Palevol 8, 179-188.View article

NEUMAYR, M. 1889. Die Stämme des Thierreiches. Wirbellose Thiere, volume 1. 603 pp. Tempsky, Vienna & Prague.

NICHOLS, D. 1962. Echinoderms. 200 pp. Hutchinson & Co., London.

NICHOLS, D. 1972. The water-vascular system in living and fossil echinoderms. Palaeontology 15, 519-538.

PARSLEY, R.L. 1970. Revision of the North American Pleurocystitidae (Rhombifera - Cystoidea). Bulletins of American Paleontology 58, 135-213.

PARSLEY, R.L. 1972. The Belemnocystitidae: solutan homeomorphs of the Anomalocystitidae. Journal of Paleontology 46, 341-347.

PARSLEY, R.L. 1982. Homalozoans, 322-323. In SPRINKLE, J. (ed.) Echinoderm Faunas from the Bromide Formation (Middle Ordovician) of Oklahoma. University of Kansas Paleontological Contributions, Monograph 1.

PARSLEY, R.L. 1990. Aristocystites, a recumbent diploporid (Echinodermata) from the Middle and Late Ordovician of Bohemia (ČSSR). Journal of Paleontology 64, 278-293.

PARSLEY, R.L. 1997. The echinoderm classes Stylophora and Homoiostelea: non Calcichordata. Paleontological Society Papers 3, 225-248.

PARSLEY, R.L. 1998. Community setting and functional morphology of Echinosphaerites infaustus (Fistuliporita: Echinodermata) from the Ordovician of Bohemia. Věstník Českého geologického ústavu 73, 253-265.

PARSLEY, R.L. 1999. The Cincta (Homostelea) as blastozoans, 369-375. In CANDIA CARNEVALI, M.D. & BONASORO, F. (eds) Echinoderm Research 1998. Balkema, Rotterdam.

PARSLEY, R.L. & CASTER, K.E. 1965. North American Soluta (Carpoidea, Echinodermata). Bulletins of American Paleontology 49(221), 109-174.

PARSLEY, R.L., ROZHNOV, S.V. & SUMRALL, C.D. 2012. Morphologic and systematic revision of the solute Maennilia estonica (Homoiostelea, Echinodermata) from the Upper Ordovician of Estonia. Journal of Paleontology 86, 462-469.View article

PARSLEY, R.L. & SUMRALL, C.S. 2007. New recumbent echinoderm genera from the Bois d’Arc Formation: Lower Devonian (Lochkovian) of Coal County, Oklahoma. Journal of Paleontology 81, 1486-1493.View article

PAUL, C.R.C. 1967. The functional morphology and mode of life of the cystoid Pleurocystites E. Billings, 1854, 105-123. In MILOT, E. (ed.) Echinoderm Biology. Symposium of the Zoological Society of London 20.

PAUL, C.R.C. 1984. British Ordovician cystoids, part 2. Palaeontolographical Society Monographs 136, 65-152.

PETERSON, K.J., ARENAS-MENA, C. & DAVIDSON, E.H. 2000. The A/P axis in echinoderm ontogeny and evolution: evidence from fossils and molecules. Evolution & Development 2, 93-101.View article

PETR, V. 1989a. Revision of morphology and ecology of Bohemura jahni Jaekel, 1903 (Ophiuroidea, Protasteridae) from Bohemian Middle Ordovician. Sborník Národního muzea v Praze 45, 1-20.

PETR, V. 1989b. Two new species of brittle-stars (Ophiuroidea, Protasteridae) from the Upper Ordovician of Bohemia. Sborník Národního muzea v Praze 45, 61-71.

PHILIP, G.M. 1979. Carpoids - echinoderms or chordates? Biological Reviews 54, 439-471.View article

PROKOP, R.J. 1984. Revision of the genus Polycrinus Jaekel, 1918 (Crinoidea, Inadunata) from the Upper Ordovician of Bohemia. Časopis pro mineralogii a geologii 29, 181-184.

PROKOP, R.J. & PETR, V. 1990. Letenocrinus longibrachialis gen. et sp. n. (Echinodermata, Paracrinoidea?) from the Bohemian Middle Ordovician. Časopis Národního muzea, Řada přirodovědná 155, 105-108.

PROKOP, R.J. & PETR, V. 1999. Echinoderms in the Bohemian Ordovician. Journal of the Czech Geological Society 44, 63-68.

PROKOP, R.J. & PETR, V. 2003. Plasiacystis mobilis gen. et sp. n., a strange “carpoid” (Echinodermata, ?Homoiostelea: Soluta) in the Bohemian Ordovician (Czech Republic). Sborník Národního muzea, Řada B - přírodní vědy 59, 151-162.

RAHMAN, I.A. & LINTZ, H. 2012. Dehmicystis globulus, an enigmatic solute (Echinodermata) from the Lower Devonian Hunsrück Slate, Germany. Paläontologische Zeitschrift 86, 59-70.View article

RAK, S. 2009. The occurrence of non-trilobite arthropods from the quartzites of Letná Formation. Český kras 35, 14-25.

REGNÉLL, G. 1945. Non-crinoid Pelmatozoa from the Paleozoic of Sweden. Meddelanden fran Lunds Geologisk-Mineralogiska Institution 108, 1-255.

ROZHNOV, S.V. 2002. Morphogenesis and evolution of crinoids and other pelmatozoan echinoderms in the Early Palaeozoic. Paleontological Journal 36 (Suppl. 6), 525-674.

ROZHNOV, S.V. 2012. The anteroposterior axis in echinoderms and displacement of the mouth in their phylogeny and ontogeny. Biology Bulletin 39, 162-171.View article

ROZHNOV, S.V. & JEFFERIES, R.P.S. 1996. A new stem-chordate solute from the Middle Ordovician of Estonia. Geobios 29, 91-109.View article

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.

SEVASTOPULO, G.D. 2008. Paleobiology of Carboniferous microcrinoids, 55-69. In AUSICH, W.I & WEBSTER, G.D. (eds) Echinoderm Paleobiology. Indiana University Press, Bloomington.

SMITH, A.B. 2005. The pre-radial history of echinoderms. Geological Journal 40, 255-280.View article

SMITH, A.B. 2008. Deuterostomes in a twist: the origins of a radical new body plan. Evolution & Development 10, 493-503.View article

SMITH, A.B. & ZAMORA, S. 2009. Rooting phylogenies of problematic fossil taxa; a case study using cinctans (stem-group echinoderms). Palaeontology 52, 803-821.View article

SPENCER, W.K. & WRIGHT, C.W. 1966. Asterozoans, 4-107. In MOORE, R.C. (ed.) Treatise on Invertebrate Paleontology. Part U, Echinodermata 3(1). Geological Society of America & University of Texas Press, Boulder & Lawrence.

SPRINKLE, J. 1973. Morphology and Evolution of Blastozoan Echinoderms. 283 pp. Museum of Comparative Zoology Harvard University, Cambridge (Massassuchetts).

SPRINKLE, J. 1974. New rhombiferan cystoids from the Middle Ordovician of Nevada. Journal of Paleontology 48, 1174-1201.

SPRINKLE, J. 1976. Biostratigraphy and paleoecology of Cambrian echinoderms from the Rocky Mountains. Brigham Young University Geology Studies 23, 61-73.

SPRINKLE, J. 1992. Radiation of Echinodermata, 375-398. In LIPPS, J.H. & SIGNOR, P.W. (eds) Origin and Early Evolution of the Metazoa. Plenum Press, New York.

SPRINKLE, J. & GUENSBURG, T.E. 2001. Growing a stalked echinoderm within the Extraxial-Axial Theory, 59-65. In BARKER, M.F. (ed.) Echinoderms 2000. Swets & Zeitlinger, Lisse.

SPRINKLE, J. & GUENSBURG, T.E. 2004. Crinozoan, blastozoan, echinozoan, asterozoan, and homalozoan echinoderms, 266-280. In WEBBY, B.D., PARIS, F., DROSER, M.L. & PERCIVAL, I.G. (eds) The Great Ordovician Biodiversification Event. Columbia University Press, New York.

SPRINKLE, J. & KIER, P.M. 1987. Phylum Echinodermata, 550-611. In BOARDMAN, R.S., CHEETHAM, A.H. & ROWELL, A.J. (eds) Fossil Invertebrates. Blackwell, London.

ŠTORCH, P., FATKA, O. & KRAFT, P. 1993. Lower Palaeozoic of the Barrandian area (Czech Republic) - a review. Coloquios de Paleontologia 45, 163-191.

STRIMPLE, H.L. 1953. A new carpoid from Oklahoma. Journal of the Washington Academy of Sciences 43, 105-106.

SUMRALL, C.D. 1997. The role of fossils in the phylogenetic reconstruction of Echinodermata. Paleontological Society Papers 3, 267-288.

SUMRALL, C.D. 2000. The biological implications of an edrioasteroid attached to a pleurocystitid rhombiferan. Journal of Paleontology 74, 67-71.View article

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, 1091-1109.

SUMRALL, C.D., SPRINKLE, J., GUENSBURG, T.E. & DATTILO, B.F. 2012. Early Ordovician mitrates and a possible solute (Echinodermata) from the western United States. Journal of Paleontology 86, 595-604.View article

TALENT, J.A. 1990. Himalayan geology again: curiouser and spuriouser! The Australian Geologist 74, 42-46.

TERMIER, H. & TERMIER, G. 1948. Les échinodermes du Paléozoique inférieur. La Revue Scientifique 3298, 613-626.

THAYER, C.W. 1975. Morphologic adaptations of benthic invertebrates to soft substrata. Journal of Marine Research 3, 177-189.

THOMAS, A.O. & LADD, H.S. 1926. Additional cystoids and crinoids from the Maquoketa Shale of Iowa. University of Iowa Studies in Natural History 11, 5-18.

THORAL, M. 1935. Contribution a l’étude paléontologique de l’Ordovicien inférieur de la Montagne Noire et révision sommaire de la faune cambrienne de la Montagne Noire. 362 pp. Imprimerie de la Charité, Montpellier.

UBAGHS, G. 1960. Le genre Lingulocystis Thoral (Echinodermata, Eocrinoidea), avec des remarques critiques sur la position systématique du genre Rhipidocystis Jaekel. Annales de Paléontologie 46, 81-116.

UBAGHS, G. 1963. Cothurnocystis Bather, Phyllocystis Thoral and an undetermined member of the order Soluta (Echinodermata, Carpoidea) in the uppermost Cambrian of Nevada. Journal of Paleontology 37, 1133-1142.

UBAGHS, G. 1966. Ophiocistioids, 174-188. In MOORE, R.C. (ed.) Treatise on Invertebrate Paleontology. Part U, Echinodermata 3(1). Geological Society of America & University of Texas Press, Boulder & Lawrence.

UBAGHS, G. 1967a. Eocrinoidea, 455-495. In MOORE, R.C. (ed.) Treatise on Invertebrate Paleontology. Part S, Echinodermata 1(2). Geological Society of America & University of Texas Press, Boulder & Lawrence.

UBAGHS, G. 1967b. Stylophora, 495-565. In MOORE, R.C. (ed.) Treatise on Invertebrate Paleontology. Part S, Echinodermata 1(2). Geological Society of America & University of Texas Press, Boulder & Lawrence.

UBAGHS, G. 1967c. Homostela, 565-581. In MOORE, R.C. (ed.) Treatise on Invertebrate Paleontology. Part S, Echinodermata 1(2). Geological Society of America & University of Texas Press, Boulder & Lawrence.

UBAGHS, G. 1970. Les échinodermes “carpoides” de l’Ordovicien inférieur de la Montagne Noire (France). 110 pp. Editions du CNRS (Cahiers de Paléontologie), Paris.

UBAGHS, G. 1979. Trois Mitrata (Echinodermata: Stylophora) nouveaux de l’Ordovicien de Tchécoslovaquie. Paläontologische Zeitschrift 53, 98-119.View article

UBAGHS, G. 1981. Réflexions sur la nature et la fonction de l’appendice articulé des “carpoides” Stylophora (Echinodermata). Annales de Paléontologie 67, 33-48.

UBAGHS, G. & ROBISON, R.A. 1985. A new homoiostelean and a new eocrinoid from the Middle Cambrian of Utah. University of Kansas Paleontological Contributions 115, 1-24.

UBAGHS, G. & ROBISON, R.A. 1988. Homalozoan echinoderms of the Wheeler Formation (Middle Cambrian) of western Utah. University of Kansas Paleontological Contributions 120, 1-17.

WAAGEN, W. & JAHN, J.J. 1899. Classe des Echinodermes. 2, Famille des Crinoides, 1-215. In BARRANDE, J. (ed.) Systeme silurien du centre de la Boheme 7. Gerhard, Leipzig & Řivnáč, Prague.

WACHSMUTH, C. & SPRINGER, F. 1881. Revision of the Palaeocrinoidea. Proceedings of the Academy of Natural Sciences of Philadelphia, 177-414.

WEBSTER, G.D. 1991. An evaluation of the V.J. Gupta echinoderm papers, 1971-1989. Journal of Paleontology 65, 1006-1008.

WETHERBY, A.G. 1881. Descriptions of new fossils from the Lower Silurian and sub-Carboniferous rocks of Kentucky. Journal of the Cincinnati Society of Natural History 4, 177-179.

WITHERS, R.B. 1933. A new genus of fossil king crabs. Proceedings of the Royal Society of Victoria 46, 18-22.

ZAMORA, S., DARROCH, S. & RAHMAN, I.A. 2013. Taphonomy and ontogeny of early pelmatozoan echinoderms: a case study of a mass-mortality assemblage of Gogia from the Cambrian of North America. Palaeogeography, Palaeoclimatology, Palaeoecology 377, 62-72.View article

ZAMORA, S., LEFEBVRE, B., ALVARO, J.J., CLAUSEN, S., ELICKI, O., FATKA, O., JELL, P., KOUCHINSKY, A., LIN, J.P., NARDIN, E., PARSLEY, R.L., ROZHNOV, S.V., SPRINKLE, J., SUMRALL, C.D., VIZCAINO, D. & SMITH, A.B. in press. Cambrian echinoderm diversity and palaeobiogeography. In HARPER, D.A.T & SERVAIS, T. (eds) Early Palaeozoic Palaeobiogeography and Palaeogeography. Geological Society of London, Memoir 38.

ZAMORA, S., ZHU, X. & LEFEBVRE, B. 2012. Palaeogeographic and evolutionary implications of new Furongian (Cambrian) echinoderms from China. Conference booklet, 14th International Echinoderm Conference, 20-24 August 2012, Brussels, 137.

ZITTEL, K.A. VON 1903. Grundzüge der Paläontologie (Paläozoologie). I. Abteilung: Invertebrata. 558 pp. Oldenbourg, Munich & Berlin.View article

ZUYKOV, M.A., TERENTIEV, S.S. & HARPER, D.A.T. 2008. New endemic brachiopod and echinoderm genera from the Upper Ordovician of the St. Petersburg region, northwestern Russia. GFF 130, 87-93.View article