Skip to main content Accessibility help

The phylogeny of the Diploporita: a polyphyletic assemblage of blastozoan echinoderms

  • Sarah L. Sheffield (a1) (a2) and Colin D. Sumrall (a2)


The phylogenetic relationships of Paleozoic blastozoan echinoderms are poorly understood and many of the traditionally ascribed groups are likely polyphyletic. Diploporitans, those blastozoans with double pore (diplopore) respiratory structures, have never been placed within a rigorous phylogenetic framework, and their highly variable morphologies suggest that they do not represent a natural clade. A maximum parsimony phylogenetic analysis, spanning a wide range of diploporitan and related taxa, indicates that diplopore-bearing blastozoans are a polyphyletic grouping and, consequently, that diplopore respiratory structures have evolved more than once within the echinoderms. Constraint analyses indicate that a single diplopore-bearing clade bearing the traditionally defined Glyptosphaeritida, Sphaeronitida, Asteroblastida is less parsimonious than multiple diplopore-bearing clades inferred by the unconstrained analysis.



Hide All
Angelin, N.P., 1878, Iconographica crinoideorum in stratis Sueciae siluricus fossilum: Stockholm, Samson & Wallin, 62p.
Ausich, W.I., Kammer, T.K., Rhenberg, E.C., and Wright, D.F., 2015, Early phylogeny of crinoids within the pelmatozoan clade: Palaentology, v. 58, p. 937952.
Barrande, J., 1846, Notice préliminaire sur le Systéme Silurien et les trilobites de Bohême: Leipzig, C. L. Hirschfeld, 97 p.
Barrande, J., 1887. Classe des échinodermes, ordre des Cystidées, in Barrande, J., Počta, F., Perner, J., Waagen, W.H., and Jahn, J., eds., Système silurien du Centre de la Bohème. Part. I : Recherches paléontologiques, ouvrage posthume de feu Joachim Barrande publié par le Docteur W. Waagen: Éditions Gerhard, v. 7, p. 1233.
Bassler, R.S., 1950, New genera of American Middle Ordovician “Cystoidea”: Washington Academy of Science, Journal, v. 40, 273277.
Bather, F.A., 1900, The Pelmatozoa-Cystoidea, in Lankester, E.R., ed., A Treatise on Zoology, Pt. 3, The Echinodermata: London, Adam and Charles Back, p. 3877.
Bernard, F., 1895, Eléments de paleontology viii: Paris, J.B. Bailliére & Fils, 612 p.
Billings, E., 1858, On the Cystidae of the lower Silurian rocks of Canada: Geological Survey of Canada Decade 3, p. 974.
Bockelie, J.F., 1979, Celticystis n. gen., a gomphocystitid cystoid from the Silurian of Sweden: Geologiska Föreningen i Stockholm Förhandlingar, v. 101, p. 157166.
Branson, E.R., and Peck, R.E., 1940, A new cystoid from the Ordovician of Oklahoma: Journal of Paleontology, v. 14, p. 8992.
Breimer, A., and Macurda, D.B., 1972, The phylogeny of the fissiculate blastoids: Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afdeling Natuurkunde, Erste Reeks 26, 390 p.
Brochu, C.A., and Sumrall, C.D., 2001, Phylogenetic nomenclature and paleontology: Journal of Paleontology, v. 75, p. 754757.
Callaway, C., 1877, On a new area of Upper Cambrian rocks in South Shropshire, with a description of new fauna: Quarterly Journal of the Geological Society of London, v. 33, p. 652672.
Chauvel, J., 1936, Note sur les Cystidées armoricaines: genre Calix et Pachycalix: Societe Geologique et Minéralogique de Bretagne, Comptes Rendus Sommaires, v. 2, p. 14.
Chauvel, J., 1941, Recherches sur les Cystoïdes et les Carpoïdes armoricaines: Mémoires de la Société Géologique et Minéralogique de Bretagne, v. 5, 286 p.
Clausen, S., 2004, New early Cambrian eocrinoids from the Iberian Chains (NE Spain) and their role in nonreefal benthic communities: Eclogae Geologicae Helveiae, v. 97, p. 371379.
Clausen, S., and Smith, A.B., 2005, Palaeoanatomy and biological affinities of a Cambrian deuterostome: Nature, v. 438, p. 351354.
Clausen, S., and Smith, A.B., 2008, Stem structure and evolution in the earliest pelmatozoan echinoderms: Journal of Paleontology, v. 82, p. 737748.
Dickson, J.A.D., 2002, Fossil echinoderms as a monitor of the Mg/Ca ratio of Phanerozoic oceans: Science, v. 298, p. 12221224.
Dickson, J.A.D., 2004, Echinoderm skeletal preservation: calcite-aragonite seas and the Mg/Ca ratio of Phanerozoic oceans: Journal of Sedimentary Research, v. 74, p. 355365.
Eichwald, E., 1840, Sur la système Silurien d l'Esthonie: St Petersburg, l'Académie de Médecine de St. Petersburg, 1840, vol. 1, p. 1222.
Eichwald, E., 1862, Asteroblastus stellatus, eine neue Sippe und Art untersilurischer Blastoideen von Pulkowa: Bulletin de la Societe Geologique de France, v. 19, p. 6264.
Foote, M., Paleozoic record of morphological diversity in blastozoan echinoderms: Proceedings of the National Academy of Sciences of the United States of America, v. 89, p. 73257329.
Frest, T.J., Strimple, H.L., and Paul, C.R.C., 2011, The North American Holocystites fauna (Echinodermata: Blastozoa: Diploporita): paleobiology and systematics: Bulletins of American Paleontology, v. 380, 141 p.
Gelman, A., 2013, Commentary: p values and statistical practice: Epidemiology, v. 24, p. 6972.
Gil Cid, M.D., and García-Rincón, J.M., 2012, Thecal (oral zone) elements in cystoids from Spain: Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 264, p. 181190.
Guensburg, T.E., and Sprinkle, J., 2007, Phylogenetic implications of the Protocrinoida: blastozoans are not ancestral to crinoids: Annales de Palentologie, v. 93, p. 277290.
Guensburg, T.E., and Sprinkle, J., 2009, Solving the mystery of crinoid ancestry: new fossil evidence of arm origin and development: Journal of Paleontology, v. 83, p. 350364.
Guensburg, T.E., Blake, D.B., Sprinkle, J., and Mooi, R., 2016, Crinoid ancestry without blastozoans: Acta Palaentologica Polonica, v. 61, p. 253266.
Gyllenhaal, J.A., 1772, Beskrifning på de så kallade Crystall-äplen och kalkbollar, såsom petreficerade Djur af Echini genus, eller dess närmaste slägtingar: Kongl Svenska Vetenskaps Academiens Handlingar, v. 33, p. 239261
Haeckel, E., 1896, Die Amphorideen und Cystoideen: Beiträge zur Morphologie und Phylogenie der Echinodermen: Festchrift zum siebenzigsten Geburtstage von Carl Gegenbaur: Leipzig, W. Engelmann, 179 p.
Hall, J., 1861, Descriptions of new species of fossils: from the investigations of the survey: Report of the Superintendent of the Geological Survey Exhibiting the Progress of the Work. Madison, Wisconsin, p. 9–52.
Jaekel, O., 1899, Stammesgeschichte der Pelmatozoen I. Thecoidea und Cystoidea: Berlin, J. Springer, 422 p.
Jaekel, O., 1918, Phylogenie und System der Pelmatozoen: Palaeontologische Zeitschrift, v. 3, p. 1128.
Jell, P.A., 2010, Late Silurian echinoderms from the Yass Basin, New South Wales—the earliest holothurian body fossil and two diploporitan cystoids (Sphaeronitidae and Holocystitidae): American Association of Petroleum Geologists Memoir, v. 39, p. 2741.
Kammer, T.W., Sumrall, C.D., Zamora, S., Ausich, W.I., and Deline, B., 2013, Oral region homologies in Paleozoic crinoids and other plesiomorphic pentaradial echinoderms: PloS one, v. 8, e77989.
Kesling, R.V., 1967, Cystoidea, in Moore, R.C., ed., Treatise on Invertebrate Paleontology, Part S, Echinodermata 1: Lawrence, Kansas, and Boulder, Colorado, University of Kansas Press and Geological Society of America, p. S85S262
Lefebvre, B., 2007, Early Palaeozoic palaeobiogeography and palaeoecology of stylophoran echinoderms: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 245, p. 156199.
Lefebvre, B., and Fatka, O., 2003, Palaeogeographical and palaeoecological aspects of the Cambro-Ordovician radiation of echinoderms in Gondwanan Africa and peri-Gondwanan Europe: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 195, p. 7397.
Lefebvre, B., Sumrall, C.D., Shroat-Lewis, R.A., Reich, M., Webster, G.D., Hunter, A.W., Nardin, E., Rozhnov, S.V., Guensberg, T.E., and Touzeau, A., 2013, Palaeobiogeography of Ordovician echinoderms: Geological Society, London, Memoirs, v. 38, p. 173198.
Miller, S.A., 1889, North American Geology and Paleontology for the use of amateurs, students, and scientists: Cincinnati, Ohio, Western Methodist Book Concern, 664 p.
Mooi, R., and David, B., 1997, Skeletal homologies of echinoderms: The Paleontological Society Papers, v. 3, 305335.
Müller, J., 1854, Über den Bau der Echinodermen: Königlich Preussische Akademie der Wissenschaften, Abhandlungen, v. 1853, p. 125220.
Nardin, E., Lefebvre, B., David, B., and Mooi, R., 2009, La diversification des échinodermes primitifs au Paléozoïque inférieur: l'exemple des blastozoaires: Comptes-Rendus PalEvol, v. 8, p. 179188.
Neumayr, M., 1889, Die Stämme des Thierreiches, Wirbellose Thiere: Vienna and Prague, 603 p.
O'Malley, C.E., Ausich, W.I., and Chin, Y., 2016, Deep echinoderm phylogeny preserved in organic molecules from Paleozoic fossils: Geology, v. 44, 379382.
Parsley, R.L., 1982, Eumorphocystis, in Sprinkle, J., ed., Echinoderm Faunas from the Bromide Formation (middle Ordovician) of Oklahoma: The University of Kansas, Paleontological Contributions, Monograph, v. 1, p. 106117.
Patterson, C., 1982, Morphological characters and homology, in Joysey, K.A., and Friday, A.E., eds., Systematics Association Special Volume 21: Problems of Phylogeny Reconstruction: New York, Academic Press, p. 2174.
Paul, C.R.C., 1968, Morphology and function of dichoporite pore-structures in cystoids: Palaeontology, v. 11, p. 697730.
Paul, C.R.C., 1971, Revision of the Holocystites Fauna (Diploporita) of North America: Fieldiana Geology, v. 24, p. 1166.
Paul, C.R.C., 1972, Morphology and function of exothecal pore-structures in cystoids: Palaeontology, v. 15, p. 128.
Paul, C.R.C., 1988, The phylogeny of the cystoids, in Paul, C.R.C., and Smith, A.B., eds., Echinoderm Phylogeny and Evolutionary Biology: Oxford, Clarendon Press, p. 199213.
Rahman, I.A., and Zamora, S., 2009, The oldest cinctan carpoid (stem-group Echinodermata), and the evolution of the water vascular system: Zoological Journal of the Linnean Society, v. 157, p. 420432.
Robison, R.A., 1965, Middle Cambrian eocrinoids from western North America: Journal of Paleontology, v. 39, p. 355364.
Rouault, M., 1851, Fossiles du terrain silurien: Bulletin de la Societe Geologique de France, v. 8, p. 358399.
Sheffield, S.L., and Sumrall, C.D., 2015, A new interpretation of the oral plating patterns of the Holocystites Fauna, in Zamora, S. and Rábano, I., eds., Progress in Echinoderm Palaeobiology: Cuademos del Museo Geominero, 19. Instituto Geológico y Minero de España, Madrid, p. 159162.
Sheffield, S.L., and Sumrall, C.D., 2017, Generic revision of the Holocystitidae of North America (Diploporita: Echinodermata) based on universal elemental homology: Journal of Paleontology, v. 91, p. 755766. doi:10.1017/jpa.2016.159
Sheffield, S.L., and Sumrall, C.D., 2019, A re-interpretation of the ambulacral system of Eumorphocystis (Blastozoa: Echinodermata) and its bearing on the evolution of early crinoids: Palaeontology v. 62, p. 163173. doi: 10.1111/pala.12396
Sheffield, S.L., Ausich, W.I., and Sumrall, C.D., 2017, Late Ordovician (Hirnantian) diploporitan fauna of Anticosti Island, Quebec, Canada: implications for evolutionary and biogeographic patterns: Canadian Journal of Earth Sciences, v. 55, p. 17.
Sprinkle, J., 1973, Blastozoan echinoderms: Cambridge, Harvard University Museum of Comparative Zoology Special Publication, 283p.
Sprinkle, J., and Bell., B.M., 1978, Paedomorphosis in edrioasteroid echinoderms: Paleobiology, v. 4, p. 8288.
Sprinkle, J., and Wahlman, G.P., 1994, New echinoderms from the Early Ordovician of west Texas: Journal of Paleontology, v. 68, p. 324388.
Sumrall, C.D., 1997, The role of fossils in the phylogenetic reconstruction of Echinodermata, in Waters, J.A., and Maples, C.G., eds., Geobiology of Echinoderms: Paleontological Society Paper, v. 3, p. 267288.
Sumrall, C.D., 2010, A model for elemental homology for the peristome and ambulacra in blastozoan echinoderms, in Harris, L.G., Böttger, S.A., Walker, C.W., and Lesser, M.P., eds., Echinoderms: Durham, London, CRC Press, p. 269276.
Sumrall, C.D. 2015. Understanding the oral area of derived stemmed echinoderms, in Zamora, S. and Rábano, I., eds. Progress in Echinoderm Palaeobiology: Cuademos del Museo Geominero, 19. Instituto Geológico y Minero de España, Madrid, p. 169173.
Sumrall, C.D., 2017. New insights concerning homology of the oral region and ambulacral system plating of pentaradial echinoderms: Journal of Paleontology, v. 91, p. 604617.
Sumrall, C.D., and Gahn, F.J., 2006, Morphological and systematic reinterpretation of two enigmatic edrioasteroids (Echinodermata) from Canada: Canadian Journal of Earth Sciences, v. 43, p. 497507.
Sumrall, C.D., and Sprinkle, J., 1995, Plating and pectinirhombs of the Ordovician rhombiferan Plethoschisma: Journal of Paleontology, v. 69, p. 772778.
Sumrall, C.D., and Waters, J.A., 2012, Universal elemental homology in glyptocystitoids, hemicosmitoids, coronoids and blastoids: steps toward echinoderm phylogenetic reconstruction in derived Blastozoa: Journal of Paleontology, v. 86, p. 956972.
Sumrall, C.D., Brett, C.E., Dexter, T.A., and Bartholomew, A., 2009, An enigmatic blastozoan echinoderm fauna from central Kentucky: Journal of Paleontology, v. 83, p. 739749.
Sumrall, C.D., Deline, B., Colmenar, J., Sheffield, S.L., and Zamora, S., 2015, New data on late Ordovician (Katian) echinoderms from Sardinia, Italy), in Zamora, S., and Rábano, I., eds., Progress in Echinoderm Palaeobiology, Cuademos del Museo Geominero, 19. Instituto Geológico y Minero de España, Madrid, p. 175178.
Swofford, D.L., 2003, PAUP* Version 4.0.b10 Phylogenetic Analysis Using Parsimony and Other Methods: Sunderland, MA, Sinauer Associates.
Templeton, A.R., 1983, Phylogenetic inference from restriction endonuclease cleavage site maps with particular reference to the evolution of humans and the apes: Evolution, v. 37, p. 221244.
Ulrich, E.O., and Kirk, E., 1921, Amecystis, a new genus of Ordovician Cystidea: Proceedings of the Biological Society of Washington, v. 34, p. 147148.
Volborth, A. von, 1846, Über die russichen Spaheroniten, eingeleitet durch einige Betrachtungen über die Arme der Cystideen: Verhandlungen der Russisch-Kaiserlichen Mineralogischen Gesellschaft zu St. Petersburg, 1845–1846, p. 161198.
Zamora, S., and Smith, A.B., 2008, A new middle Cambrian stem-group echinoderm from Spain: paleobiological implications of a highly asymmetric cinctan: Acta Palaeontologica Polonica, v. 53, p. 207221.
Zamora, S., and Rahman, I.A., 2014, Deciphering the early evolution of echinoderms with Cambrian fossils: Palaentology, v. 57, p. 11051119.
Zamora, S., Lefebvre, B., Àlvaro, J.J., Clausen, S., Elicki, O., Fatka, O., Jell, P., Kouchinsky, A., Lin, J.-P., Nardin, E., Parsley, R., Rozhnov, S.V., Sprinkle, J., Sumrall, C.D., Vizcaino, D., and Smith, A.B., 2013, Cambrian echinoderm diversity and palaeobiogeography: Geological Society, London, Memoirs, v. 38, p. 157171.
Zamora, S., Sumrall, C.D., Zhu, X-J., and Lefebvre, B., 2016, A new stemmed echinoderm from the Furongian of China and the origin of Glyptocystitida (Blastozoa, Echinodermata): Geological Magazine, v. 154, p. 111.

Related content

Powered by UNSILO

The phylogeny of the Diploporita: a polyphyletic assemblage of blastozoan echinoderms

  • Sarah L. Sheffield (a1) (a2) and Colin D. Sumrall (a2)


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.