Skip to main content
SpringerLink
Log in

New Siluro-Devonian Anomalocystitids (Echinodermata, Stylophora) from Bolivia and Canada, and a Reevaluation of Skeletal Homologies in Mitrates

  • Published:
Paleontological Journal Aims and scope Submit manuscript

Abstract

Two new mitrate stylophorans are described from the upper part of the Santa Rosa Formation (Lochkovian) in the Central Subandean Zone of Bolivia and from the Jupiter Formation (Cybèle Member, late Llandovery) of Anticosti Island (Canada). Both stylophorans are identified as allanicytidiine anomalocystitids. The new Bolivian mitrate, Perikefalea racheboeufi gen. and sp. nov. is characterized by a strongly reduced number of plates (five) on its upper thecal surface and a strong spike on its posterior median marginal. The morphology of P. racheboeufi suggests relatively strong affinities with Occultocystis, from the Lower Devonian Talacasto Formation of Argentina. Although incompletely preserved, the Anticosti mitrate, Perikefalea? cybeleae sp. nov., represents the first stylophoran described in the Silurian of North America and the first report of post-Ordovician Allanicytidiinae outside of (peri) Gondwanan regions. Skeletal homologies in mitrates (marginals, infracentrals, posterior spines, adorals, supracentrals) are reevaluated, so as to describe the plate pattern of Perikefalea. This revision was prompted by new fossil evidence from the Furongian–Tremadocian interval suggesting that (1) mitrates derive from “Phyllocystis” jingxiensis-like cornutes with wide marginals, a posteriorly open marginal frame (loss of M5 and \({\text{M}}_{5}^{'}\)), a zygal plate in central position, and confluent, sutured M3 and G; (2) anomalocystitids derive from Mitrocystites-like mitrocystitidans by modification of two marginals (digital and glossal) into posterior spines. Two distinct terminologies are tentatively proposed for supracentrals in Allanicytidiinae and Anomalocystitinae, because the highly standardized plate patterns of their Siluro-Devonian representatives were possibly acquired independently in the two clades.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Log in via an institution

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Institutional subscriptions

Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
Fig. 13.
Fig. 14.
Fig. 15.
Fig. 16.
Fig. 17.
Fig. 18.
Fig. 19.
Fig. 20.
Fig. 21.
Fig. 22.
Fig. 23.

Similar content being viewed by others

REFERENCES

  1. Ausich, W.I. and Copper, P., The Crinoidea of Anticosti Island, Québec (Late Ordovician to Early Silurian), Palaeontogr. Can., 2010, vol. 29, pp. 1–157.

    Google Scholar 

  2. Ausich, W.I. and Cournoyer, M.E., New taxa and revised stratigraphic distribution of the crinoid fauna from Anticosti Island, Québec, Canada (Late Ordovician–Early Silurian), J. Paleontol., 2019, vol. 93, pp. 1137–1158.

    Article  Google Scholar 

  3. Barnes, C.R., Stratigraphy and paleontology of the Ordovician-Silurian boundary interval, Anticosti Island, Québec, Canada, Bull. Br. Mus. (Nat. Hist.), Geol., 1988, vol. 43, pp. 195–219.

    Google Scholar 

  4. Bassler, R.S., The stratigraphy of the Central Basin of Tennessee, Bull. – Ten., Div. Geol., 1932, vol. 38, pp. 1–268.

    Google Scholar 

  5. Bassler, R.S., Pelmatozoa Palaeozoica (Generum et Genotyporum Index et Bibliographia), in Fossilium Catalogus, I, Animalia, pars 83, Quenstedt, W., Ed., Gravenhage: Verlag Naturwiss., 1938, pp. 1–194.

    Google Scholar 

  6. Bather, F.A., Caradocian Cystidea from Girvan, Trans. – R. Soc. Edinburgh, 1913, vol. 49, pp. 359–529.

    Article  Google Scholar 

  7. Bolton, T.E., Ordovician and Silurian formations of Anticosti Island, Quebec, Pap. - Geol. Surv. Can., 1961, vol. 71–19, pp. 1–45.

  8. Botting, J.P., Muir, L.A., and Lefebvre, B., Echinoderm diversity and environmental distribution in the Ordovician of the Builth Inlier, Wales, Palaios, 2013, vol. 28, pp. 293–304.

    Article  Google Scholar 

  9. Boucot, A.J., Devonian biogeography: an update, in Devonian of the World, McMillan, N.J., Embry, A.F., and Glass, D.J., Eds., Proc. Sec. Int. Symp. Dev. Syst. Calgary, 1988, vol. 3, pp. 211–227.

  10. Boucot, A.J. and Rachebœuf, P.R., Biogeographic summary of the Malvinokaffric Realm Silurian and Devonian fossils, in Fosiles y Facies de Bolivia, vol. II: Invertebrados y Paleobotanica, Suarez-Soruco, R., Ed., Rev. Técnica YPFB, 1993, vol. 13–14, pp. 71–75.

  11. Bruguière, J.G., 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, 1791, pp. 1–180.

    Google Scholar 

  12. Caster, K.E., Concerning Enoploura of the Upper Ordovician and its relation to other carpoid Echinodermata, Bull. Am. Paleontol., 1952, vol. 34, pp. 1–47.

    Google Scholar 

  13. Caster, K.E., A new carpoid from the Parana Devonian, An. Acad. Bras. Cienc., 1954a, vol. 26, pp. 123–147.

    Google Scholar 

  14. Caster, K.E., A Devonian placocystoid echinoderm from Parana, Brazil, in Paleontologia do Paraná, Lange, F.W., Ed., Caritiba: Com. Comemor. Centenario Paraná, 1954b, pp. 137–148.

  15. Caster, K.E., A new Silurian carpoid echinoderm from Tasmania and a revision of the Allanicytidiidae, Alcheringa, 1983, vol. 7, pp. 321–335.

    Article  Google Scholar 

  16. Caster, K.E. and Eaton, J.R., Microstructure of the plates in the carpoid echinoderm Paranacystis, J. Paleontol., 1956, vol. 30, pp. 611–614.

    Google Scholar 

  17. Chauvel, J., Recherches sur les cystoïdes et les carpoïdes armoricains, Mem. Soc. Geol. Mineral. Bretagne, 1941, vol. 5, pp. 1–286.

    Google Scholar 

  18. Chauvel, J., Etude critique de quelques échinodermes stylophores du Massif armoricain, Bull. Soc. Geol. Mineral. Bretagne (C), 1981, vol. 13, pp. 67–101.

    Google Scholar 

  19. Chen, G.Y. and Han, N.R., New materials of Stylophora from the Upper Cambrian of the Jingxi area, Guangxi, South China, Acta Palaeontol. Sin., 2013, vol. 52, pp. 288–293.

    Google Scholar 

  20. Cocks, L.R.M. and Copper, P., The Ordovician-Silurian boundary at the eastern end of Anticosti Island, Can. J. Earth Sci., 1981, vol. 18, pp. 1029–1034.

    Article  Google Scholar 

  21. Copper, P., Upper Ordovician and Lower Silurian reefs of Anticosti Island, Quebec, Can. Soc. Petrol. Geol. Mem, 1989, vol. 13, pp. 271–276.

    Google Scholar 

  22. Copper, P., Reefs during the multiple crises towards the Ordovician-Silurian boundary: Anticosti Island, eastern Canada, and worldwide, Can. J. Earth Sci., 2001, vol. 38, pp. 153–171.

    Article  Google Scholar 

  23. Copper, P., Jin, J., and Desrochers, A., The Ordovician-Silurian boundary (late Katian–Hirnantian) of western Anticosti Island: revised stratigraphy and benthic megafauna correlations, Stratigraphy, 2013, vol. 19, pp. 213–227.

    Google Scholar 

  24. Craske, A.J. and Jefferies, R.P.S., A new mitrate from the Upper Ordovician of Norway and a new approach to subdividing a plesion, Palaeontology, 1989, vol. 32, pp. 69–99.

    Google Scholar 

  25. Cripps, A.P., A new stem-craniate from the Ordovician of Morocco and the search for the sister group of the Craniata, Zool. J. Linn. Soc., 1990, vol. 100, pp. 27–71.

    Article  Google Scholar 

  26. Cripps, A.P., A cladistic analysis of the cornutes (stem chordates), Zool. J. Linn. Soc., 1991, vol. 102, pp. 333–366.

    Article  Google Scholar 

  27. David, B., Lefebvre, B., Mooi, R., and Parsley, R., Are homalozoans echinoderms? An answer from the extraxial-axial theory, Paleobiology, 2000, vol. 26, pp. 529–555.

    Article  Google Scholar 

  28. Dehm, R., Cystoideen aus dem rheinischen Unterdevon, N. Jb. Miner. Geol. Paläont. Abt. A, 1932, vol. 69, pp. 63–93.

    Google Scholar 

  29. Dehm, R., Untersuchungen an Cystoideen des rheinischen Unterdevons, Sitzungsber. - Bayer. Akad. Wiss., Math.-Naturwiss. Kl. (Abteilung), 1934, pp. 19–34.

    Google Scholar 

  30. De Koninck, M.L., Sur quelques échinodermes remarquables des terrains paléozoïques, Bull. Acad. R. Belg., 1869, vol. 28, pp. 544–552.

    Google Scholar 

  31. Derstler, K., Biogeography of the stylophoran carpoids (Echinodermata), in Historical Biogeography, Plate Tectonics, and the Changing Environment, Gray, J. and Boucot, A.J., Eds., Corvallis: Oregon State Univ. Press, 1979, pp. 91–104.

    Google Scholar 

  32. Desrochers, A., Farley, C., Achab, A., and Asselin, E., A high-resolution stratigraphic model to resolve the longstanding issues relative to the correlation and interpretation of the O/S boundary on Anticosti Island, Canada, in Palaeozoic Climates—International Congress: Closing meeting of the International Geoscience Programme 503 “Ordovician Palaeogeography and Palaeoclimate, Abstracts, Kröger, B. and Servais T., Eds., Lille: Univ. Lille, 2008, p. 32.

  33. Desrochers, A., Farley, C., Achab, A., Asselin, E., and Riva, J.F., A far-field record of the end Ordovician glaciations: the Ellis Bay Formation, Anticosti Island, Eastern Canada, Palaeogeogr., Palaeoclimatol., Palaeoecol., 2010, vol. 296, pp. 248–263.

    Article  Google Scholar 

  34. Di Pasquo, M.M. and Noetinger, S., First record of Early Devonian (Lochkovian) flora from the Santa Rosa Formation—Alarache, southern Bolivia, Geol. Acta, 2008, vol. 6, 191–210.

    Google Scholar 

  35. Ewin, T.A.M., Martin, M., Isotalo, P., and Zamora, S., New rhenopyrgid edrioasteroids (Echinodermata) and their implications for taxonomy, functional morphology, and paleoecology, J. Paleontol., 2020, vol. 94, pp. 115–130.

    Article  Google Scholar 

  36. Gaillard, C. and Rachebœuf, P.R., Trace fossils from nearshore to offshore environments: Lower Devonian of Bolivia, J. Paleontol., 2006, vol. 80, pp. 1205–1226.

    Article  Google Scholar 

  37. Gill, E.D. and Caster, K.E., Carpoid echinoderms from the Silurian and Devonian of Australia, Bull. Am. Paleontol., 1960, vol. 41, pp. 5–71.

    Google Scholar 

  38. Gislen, T., Affinities between the Echinodermata, Enteropneusta and Chordonia, Zool. Bidr. Uppsala, 1930, vol. 12, pp. 199–304.

    Google Scholar 

  39. Guensburg, T.E. and Sprinkle, J., Ecologic radiation of Cambro-Ordovician echinoderms, in Ecology of the Cambrian Radiation, Zhuravlev, A.Y. and Riding, R., Eds., 2000, N.Y.: Columbia Univ. Press, pp. 428–444.

    Google Scholar 

  40. Haeckel, E., Die Amphorideen und Cystoideen. Beitrage zur Morphologie und Phylogenie der Echinodermen, Festschr. Siebenziegsten Geburstage von C. Gegenbaur, 1896, vol. 1, pp. 32–45.

    Google Scholar 

  41. Hall, J., Natural History of New York, Paleontology, Vol. III: Containing Descriptions and Figures of the Organic Remains of the Lower Helderburg Group and the Oriskany Sandstone (Part I, Text), Albany: Van Benthuysen, 1859, pp. 1–532.

    Google Scholar 

  42. Han, N.R. and Chen, G.Y., New stylophorans (Echinodermata) from the Upper Cambrian of Guangxi, South China, Sci. China, Ser. D: Earth Sci, 2008, vol. 51, pp. 181–186.

    Article  Google Scholar 

  43. Haude, R., Kaum bekannte und seltene Echinodermen aus dem Mitteldevon des rechtsrheinischen Schiefergebirges, Der Aufschluss, 1983, vol. 34, pp. 101–110.

    Google Scholar 

  44. Haude, R., Echinodermen aus dem Unter-Devon der argentinischen Präkordillere, N. Jb. Geol. Paläont. Abh., 1995, vol. 197, pp. 37–86.

    Google Scholar 

  45. Isaacson, P.E. and Díaz-Martinez, E., Evidence for a middle-late Paleozoic foreland basin and significant paleolatitudinal shift, Central Andes, Mem. – Am. Assoc. Pet. Geol., 1995, vol. 62, pp. 231–249.

    Google Scholar 

  46. Jaekel, O., Phylogenie und System der Pelmatozoen, Paläont. Z., 1918, vol. 3, pp. 1–128.

    Google Scholar 

  47. Janvier, P. and Maisey, J.G., The Devonian vertebrates of South America and their biogeographical relationships, in Morphology, Phylogeny and Paleobiogeography of Fossil Fishes, Elliott, D.K., Maisey, J.G., Yu, X. and Miao, D., Eds., Munich: Verlag Dr. Friedrich Pfeil, 2010, pp. 431–460.

    Google Scholar 

  48. Jefferies, R.P.S., The subphylum Calcichordata (Jefferies 1967) primitive fossil chordates with echinoderm affinities, Bull. Br. Mus. (Nat. Hist.), Geol., 1968, vol. 16, pp. 243–339.

    Google Scholar 

  49. Jefferies, R.P.S. Ceratocystis perneri Jaekel—A Middle Cambrian chordate with echinoderm affinities, Palaeontology, 1969, vol. 12, pp. 494–535.

    Google Scholar 

  50. Jefferies, R.P.S., The Ordovician fossil Lagynocystis pyramidalis (Barrande) and the ancestry of amphioxus, Philos. Trans. R. Soc., B, 1973, vol. 265, pp. 409–469.

  51. Jefferies, R.P.S, Locomotion, shape, ornament, and external ontogeny in some mitrate calcichordates, J. Vert. Paleontol., 1984, vol. 4, pp. 292–319.

    Article  Google Scholar 

  52. Jefferies, R.P.S., The chordates—a preliminary note, in The Arenig Series in South Wales: Stratigraphy and Palaeontology, Fortey, R.A and Owens, R.M., Eds., Bull. Br. Mus. (Nat. Hist.), Geol., 1987, vol. 41, pp. 285–290.

  53. Jefferies, R.P.S. and Lewis, D.N., The English Silurian fossil Placocystites forbesianus and the ancestry of the vertebrates, Philos. Trans. R. Soc., B, 1978, vol. 282, pp. 205–323.

  54. Jefferies, R.P.S. and Prokop, R.J., A new calcichordate from the Ordovician of Bohemia and its anatomy, adaptations and relationships, Biol. J. Linn. Soc., 1972, vol. 4, pp. 69–115.

    Article  Google Scholar 

  55. Jell, P., Placocystella in the Early Devonian (Lochkovian) of central Victoria, Alcheringa, 2013, vol. 37, pp. 567–569.

    Article  Google Scholar 

  56. Jin, J. and Copper, P., Response of brachiopod communities to environmental change during the Late Ordovician mass extinction interval, Anticosti Island, Eastern Canada, Fossils Strata, 2008, vol. 54, pp. 41–51.

    Google Scholar 

  57. Kirk, E., The structure and relationships of certain eleutherozoic Pelmatozoa, Proc. U. S. Natl. Mus., 1911, vol. 41, pp. 1–137.

    Article  Google Scholar 

  58. Klein, J.T., Naturalis Dispositio Echinodermatum. Accessit Lucubratiuncula de Aculeis Echinorum Marinorum, cum Spicilegio de Belemnitis, Dantzig: Schreiber, 1734, pp. 1–79.

  59. Kolata, D.R. and Guensburg, T.E., Diamphidiocystis, a new mitrate “carpoid” from the Cincinnatian (Upper Ordovician) Maquoketa Group in southern Illinois, J. Paleontol., 1979, vol. 53, pp. 1121–1135.

    Google Scholar 

  60. Kolata, D.R. and Jollie, M., Anomalocystitid mitrates (Stylophora-Echinodermata) from the Champlainian (Middle Ordovician) Guttenberg Formation of the Upper Mississipi Valley Region, J. Paleontol., 1982, vol. 56, pp. 631–653.

    Google Scholar 

  61. Kolata, D.R., Frest, T.J., and Mapes, R.H., The youngest carpoid: occurrence, affinities, and life mode of a Pennsylvanian (Morrowan) mitrate from Oklahoma, J. Paleontol., 1991, vol. 65, pp. 844–855.

    Article  Google Scholar 

  62. Lee, S.B., Lefebvre, B., and Choi, D.K., Uppermost Cambrian cornutes (Echinodermata, Stylophora) from the Taebaeksan Basin, Korea, J. Paleontol., 2005, vol. 79, pp. 139–151.

    Article  Google Scholar 

  63. Lefebvre, B., Stylophores (Cornuta, Mitrata): Situation au sein du Phylum des Echinodermes et Phylogenèse, Lyon: unpubl. PhD Thesis, 1999, pp. 1–630.

  64. Lefebvre, B., Les échinodermes stylophores du Massif armoricain, Bull. Soc. Sci. Nat. Ouest Fr., 2000a, vol. 22, pp. 101–122.

    Google Scholar 

  65. Lefebvre, B., A new mitrate (Echinodermata, Stylophora) from the Tremadoc of Shropshire (England) and the origin of the Mitrocystitida, J. Paleontol., 2000b, vol. 74, pp. 890–905.

    Article  Google Scholar 

  66. Lefebvre, B., Homologies in Stylophora: a test of the “calcichordate theory”, Geobios, 2000c, vol. 33, pp. 359–364.

    Article  Google Scholar 

  67. Lefebvre, B., A critical comment on “ankyroids”, Geobios, 2001, vol. 34, pp. 597–627.

    Article  Google Scholar 

  68. Lefebvre, B., Stephen J. Gould, les mitrates et les monstres, C. R. Palevol., 2003a, vol. 2, pp. 509–522.

    Article  Google Scholar 

  69. Lefebvre, B., Functional morphology of stylophoran echinoderms, Palaeontology, 2003b, vol. 46, pp. 511–555.

    Article  Google Scholar 

  70. Lefebvre, B., Stylophoran supertrees revisited, Acta Palaeontol. Pol., 2005, vol. 50, pp. 477–486.

    Google Scholar 

  71. Lefebvre, B., Early Palaeozoic palaeobiogeography and palaeoecology of stylophoran echinoderms, Palaeogeogr., Palaeoclimatol., Palaeoecol., 2007a, vol. 245, pp. 156–199.

    Article  Google Scholar 

  72. Lefebvre, B., Changes in Late Ordovician (pre-Hirnantian) echinoderm assemblages: palaeoenvironmental and palaeobiogeographical implications, Acta Palaeontol. Sin., 2007b, vol. 46 (suppl.), pp. 241–248.

    Google Scholar 

  73. Lefebvre, B. and Botting, J.P., First report of the mitrate Peltocystis cornuta Thoral (Echinodermata, Stylophora) in the Lower Ordovician of central Anti-Atlas (Morocco), Ann. Paléontol., 2007, vol. 93, pp. 183–198.

    Article  Google Scholar 

  74. Lefebvre, B. and Fatka, O., Palaeogeographical and palaeoecological aspects of the Cambro-Ordovician radiation of echinoderms in Gondwanan Africa and peri-Gondwanan Europe, Palaeogeogr., Palaeoclimatol., Palaeoecol., 2003, vol., 195, pp. 73–97.

    Article  Google Scholar 

  75. Lefebvre, B. and Gutiérrez-Marco, J.C., New Ordovician mitrocystitidan mitrates (Echinodermata, Stylophora) from the Central-Iberian zone (Spain), N. Jb. Geol. Paläontol. Abh., 2003, vol. 227, pp. 39–61.

    Google Scholar 

  76. Lefebvre, B. and Rachebœuf, P., First report of mitrate stylophorans (Echinodermata) in the Lower Devonian of Bolivia, in 4th European Meeting on the Palaeontology and Stratigraphy of Latin America, Díaz-Martinez, E. and Ríbano, I., Eds., Cuad. Mus. Geomin., 2007, vol. 8, pp. 239–244.

  77. Lefebvre, B. and Vizcaïno, D., New Ordovician cornutes (Echinodermata, Stylophora) from Montagne Noire and Brittany (France) and a revision of the Order Cornuta Jaekel 1901, Geobios, 1999, vol. 32, pp. 421–458.

    Article  Google Scholar 

  78. Lefebvre, B., Rachebœuf, P., and David, B., Homologies in stylophoran echinoderms, in Echinoderms: San Francisco, Mooi, R. and Telford, M., Eds., Rotterdam: Balkema, 1998, pp. 103–109.

    Google Scholar 

  79. Lefebvre, B., Eble, J.J., Navarro, N., and David, B., Diversification of atypical Paleozoic echinoderms: a quantitative survey of patterns of stylophoran disparity, diversity, and geography, Paleobiology, 2006, vol. 32, pp. 483–510.

    Article  Google Scholar 

  80. Lefebvre, B., Régnault, S., Lardeux, H., Kundura, J.P., and Roussel, P., New Ordovician mitrates (Echinodermata, Stylophora) from the Ancenis Basin (South Armorican Domain, France): palaeogeographic and palaeoenvironmental implications, in Echinoderms: Durham, Harris, L.G., Böttger, S.A., Walker, C.W., and Lesser, M.P., Eds., Leiden: Balkema, 2010, 37–44.

    Google Scholar 

  81. 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., and Touzeau, A., Palaeobiogeography of Ordovician echinoderms, in Early Palaeozoic Biogeography and Palaeogeography, Harper, D.A.T and Servais, T., Eds, Geol. Soc. London Mem., 2013, vol. 38, pp. 173–198.

    Google Scholar 

  82. Lefebvre, B., Allaire, N., Guensburg, T.E., Hunter, A.W., Kouraïss, K., Martin, E.L.O., Nardin, E., Noailles, F., Pittet, B., Sumrall, C.D., and Zamora, S., Palaeoecological aspects of the diversification of echinoderms in the Lower Ordovician of central Anti-Atlas, Morocco, Palaeogeogr., Palaeoclimatol., Palaeoecol., 2016, vol. 460, pp. 97–121.

    Article  Google Scholar 

  83. Lefebvre, B., Guensburg, T.E., Martin, E.L.O., Mooi, R., Nardin, E., Nohejlová, M., Saleh, F., Kouraïss, K., El Hariri, K., and David, B., Exceptionally preserved soft parts in fossils from the Lower Ordovician of Morocco clarify stylophoran affinities within basal deuterostomes, Geobios, 2019, vol. 52, pp. 27–36.

    Article  Google Scholar 

  84. Lindström, G., List of the Fossil Faunas of Sweden, Part 2, Upper Silurian, Stockholm: Norstedt & Söner, 1888, pp. 1–29.

    Google Scholar 

  85. Long, D.G.F. and Copper, P., Stratigraphy of the Upper Ordovician upper Vauréal and Ellis Bay formations, eastern Anticosti Island, Québec, Can.J. Earth Sci., 1987, vol. 24, pp. 1807–1820.

    Article  Google Scholar 

  86. Marti Mus, M., The Ordovician cornute Flabellicystis rushtoni n. gen. n. sp. (Stylophora, Echinodermata) and its phylogenetic position within the group Cornuta, Paläontol. Z., 2002, vol. 76, pp. 99–116.

    Google Scholar 

  87. Martin, E.L.O., Lefebvre, B., and Vaucher, R., Taphonomy of a stylophoran-dominated assemblage in the Lower Ordovician of Zagora area (central Anti-Atlas, Morocco), in Progress in Echinoderm Palaeobiology, Zamora, S. and Rabano, I., Eds., Cuadernos del Museo Geominero, 2015, vol. 19, pp. 95–100.

  88. McDermott, P.D. and Paul, C.R.C., Ateleocystites? lansae sp. nov. (Mitrata, Anomalocystitidae) from the Upper Ordovician of South Wales, Geol. Mag., 2017, vol. 52, pp. 1–13.

    Google Scholar 

  89. Noailles, F. and Lefebvre, B., A New Mitrate Stylophoran from the Lower Ordovician of the Central Anti-Atlas (Morocco) and the Origin of the Anomalocystitids, Conference booklet, 14th International Echinoderm Conference, Brussels, 2012.

  90. Parsley, R.L., Homalozoans, in Echinoderm Faunas from the Bromide Formation (Middle Ordovician) of Oklahoma, Sprinkle, J., Ed., Univ. Kans. Paleontol. Contrib., Pap., 1982, vol. 1, pp. 322–323.

  91. Parsley, R.L., Feeding and respiratory strategies in Stylophora, in Echinoderm Phylogeny and Evolutionary Biology, Paul, C.R.C. and Smith, A.B., Eds., 1988, Oxford: Clarendon Press, pp. 347–361.

    Google Scholar 

  92. Parsley, R.L., Review of selected North American mitrate stylophorans (Homalozoa: Echinodermata), Bull. Am. Paleontol., 1991, vol. 100, pp. 5–57.

    Google Scholar 

  93. Parsley, R.L., The echinoderm classes Stylophora and Homoiostelea: non Calcichordata, Paleontol. Soc. Pap., 1997, vol. 3, pp. 225–248.

    Google Scholar 

  94. Parsley, R.L. and Gutiérrez-Marco, J.C., Stylophorans in middle Arenig shallow water siliciclastics: Vizcainocarpus from the Imfout syncline in Morocco’s western Meseta, Bull. Geosci., 2005, vol. 80, pp. 185–192.

    Google Scholar 

  95. Parsley, R.L. and Sumrall, C.D., New recumbent echinoderm genera from the Bois d’Arc Formation: Lower Devonian (Lochkovian) of Coal County, Oklahoma, J. Paleontol., 2007, vol. 81, pp. 1486–1493.

    Article  Google Scholar 

  96. Peng, S.C., Yang, X.F., Liu, Y., Zhu, X.J., Sun, H.J., Zamora, S., Mao, Y.Y., and Zhang, Y.C., Fulu Biota, a new exceptionally-preserved Cambrian fossil assemblage from the Longha Formation in southeastern Yunnan, Palaeoworld (in press).

  97. Penn-Clarke, C.R., The Malvinokaffric Realm in the Early-Middle Devonian of South Africa, J. Afr. Earth Sci., 2019, vol. 158 (103549), pp. 1–19.

    Article  Google Scholar 

  98. Petryk, A.A., Stratigraphy, sedimentology, and paleogeography of the Upper Ordovician-Lower Silurian of Anticosti Island, Québec, in Field Meeting Anticosti-Gaspé, Québec, Lespérance, P.J., Ed., Montréal: Subcomm. Silurian Stratigr., Ordovician-Silurian Boundary Working Group., 1981, vol. 2, pp. 10–39.

    Google Scholar 

  99. Philip, G.M, Notocarpos garratti gen. et sp. nov., a new Silurian mitrate carpoid from Victoria, Alcheringa, 1981, vol. 3, pp. 29–38.

    Article  Google Scholar 

  100. Prokop, R.J., Dalejocystis n. gen. the first representative of the Carpoidea in the Devonian of Bohemia, J. Paleontol., 1963, vol. 37, pp. 648–650.

    Google Scholar 

  101. Rachebœuf, P.R., Le Hérissé, A., Paris, F., Babin, C., Guillocheau, F., Truyols-Massoni, M. and Súarez-Soruco, R., Le Dévonien de Bolivie: biostratigraphie et chronostratigraphie, Acad. Sci., Paris, C. R.,1993, vol. 317, pp. 795–802.

  102. Rahman, I.A., Zamora, S., and Geyer, G., The oldest stylophoran echinoderm: a new Ceratocystis from the Middle Cambrian of Germany, Paläontol. Z., 2010, vol. 84, pp. 227–237.

    Google Scholar 

  103. Régnault, S. and Chauvel, J., Découverte d’un échinoderme carpoïde (Stylophora – Mitrata) dans le Dévonien inférieur du Maroc, Geobios, 1987, vol. 20, pp. 669–674.

    Article  Google Scholar 

  104. Regnell, G., Non-crinoid Pelmatozoa from the Paleozoic of Sweden, A taxonomic study, Medd. Lunds Geol.-Mineral. Inst., 1945, vol. 108, pp. 1–255.

    Google Scholar 

  105. Regnell, G., The Lower Palaeozoic echinoderm faunas of the British Isles and Balto-Scandia, Palaeontology, 1960, vol. 2, pp. 161–179.

    Google Scholar 

  106. Reich, M., Lefebvre, B., and Kutscher, M., Mitrates (Echinodermata: Stylophora) from the Silurian of Gotland, Sweden, in The Seventh Baltic Stratigraphical Conference. Abstracts and Field Guide, Hints, O., Ainsaar, L., Männik, P. and Meidla, T., Eds., Tallinn: Geol. Soc. Estonia, 2008.

  107. Reid, M., Bordy, E.M., and Taylor, W., Taphonomy and sedimentology of an echinoderm obrution bed in the Lower Devonian Voorstehoek Formation (Bokkeveld Group, Cape Supergroup) of South Africa, J. Afr. Earth Sci., 2015, vol. 110, pp. 135–149.

    Article  Google Scholar 

  108. Rennie, J.V.L., On Placocystella, a new genus of cystids from the Lower Devonian of South Africa, Ann. S. Afr. Mus., 1936, vol. 31, pp. 269–275.

    Google Scholar 

  109. Richardson, J., Report of the year 1856, Canadian Surv. Report Progr. 1853-54-55-56, 1857, pp. 191–245.

  110. Rozhnov, S.V., New representatives of the class Stylophora (echinoderms), Paleontol. J., 1990, vol. 4, pp. 27–38.

    Google Scholar 

  111. Rozhnov, S.V. and Parsley, R.L., A new cornute (Homalozoa: Echinodermata) from the uppermost middle Cambrian (Stage 3, Furongian) from northern Iran: its systematics and functional morphology, Paleontol. J., 2017, vol. 51, pp. 500–509.

    Article  Google Scholar 

  112. Rozhnov, S.V., Mirantsev, G.V., and Terentiev, S.S., First report of the mitrate Lagynocystis (Echinodermata: Stylophora) in the Ordovician of the Baltic paleobasin, Zoosymposia, 2019, vol. 15, pp. 115–122.

    Article  Google Scholar 

  113. Ruta, M., First record of a paranacystid mitrate from the Bokkeveld Group of South Africa, Palaeontol. Afr., 1997a, vol. 34, pp. 15–25.

    Google Scholar 

  114. Ruta, M., Redescription of the Australian mitrate Victoriacystis with comments on its functional morphology, Alcheringa, 1997b, vol. 21, pp. 81–101.

    Article  Google Scholar 

  115. Ruta, M., First record of the mitrate Barrandeocarpus from England, Paläontol. Z, 1997c, vol. 71, pp. 97–105.

    Google Scholar 

  116. Ruta, M., A new mitrate from the Lower Ordovician of southern France, Palaeontology, 1997d, vol. 450, pp. 363–383.

    Google Scholar 

  117. Ruta, M., A cladistic analysis of the anomalocystitid mitrates, Zool. J. Linn. Soc., 1999a, vol. 127, pp. 345–421.

    Article  Google Scholar 

  118. Ruta, M., A new stylophoran echinoderm, Juliaecarpus milnerorum, from the Late Ordovician Upper Ktaoua Formation of Morocco, Bull. Br. Mus. (Nat. Hist.), Geol., Geol., 1999b, vol. 55, pp. 47–79.

    Google Scholar 

  119. Ruta, M., A species-level supertree for stylophoran echinoderms. Acta Palaeontol. Pol., 2003, vol. 48, pp. 559–568.

    Google Scholar 

  120. Ruta, M. and Bartels, C., A redescription of the anomalocystitid mitrate Rhenocystis latipedunculata from the Lower Devonian of Germany, Palaeontology, 1998, vol. 41, pp. 771–806.

    Google Scholar 

  121. Ruta, M. and Jell, P., Protocytidium gen. nov., a new anomalocystitid mitrate from the Victorian Latest Ordovician and evolution of the Allanicytidiidae, Mem. Queensl. Mus., 1999a, vol. 43, pp. 353–376.

    Google Scholar 

  122. Ruta, M. and Jell, P., Adoketocarpus gen. nov., a mitrate from the Ludlovian Kilmore Siltstone and Lochkovian Humevale Formation of central Victoria, Mem. Queensl. Mus., 1999b, vol. 43, pp. 377–398.

    Google Scholar 

  123. Ruta, M. and Jell, P., Two new anomalocystitid mitrates from the Lower Devonian Humevale Formation of central Victoria, Mem. Queensl. Mus., 1999c, vol. 43, pp. 399–422.

    Google Scholar 

  124. Ruta, M. and Jell, P., A note on Victoriacystis wilkinsi (Anomalocystitidae: Mitrata) from the Upper Silurian of Victoria, Mem. Queensl. Mus., 1999d, vol. 43, pp. 423–430.

    Google Scholar 

  125. Ruta, M. and Jell, P., Revision of Silurian and Devonian Allanicytidiidae (Anomalocystitida: Mitrata) from southeastern Australia, Tasmania and New-Zealand, Mem. Queensl. Mus., 1999e, vol. 43, pp. 431–451.

    Google Scholar 

  126. Ruta, M. and Theron, J.N., Two Devonian mitrates from South Africa, Palaeontology, 1997, vol. 40, pp. 201–243.

    Google Scholar 

  127. Scheffler, S.M., Horodyski, R.S., and Bosetti, E.P., Morphology, palaeoecology and taphonomy of the Devonian mitrate Placocystella langei from Paraná Basin, Brazil, Alcheringa, 2019, vol. 43, pp. 228–240.

    Article  Google Scholar 

  128. Schuchert, C. and Twenhofel, W.H., Ordovicic-Siluric section of the Mingan and Anticosti islands, Gulf of St. Lawrence, Geol. Soc. Am. Bull., 1910, vol. 21, pp. 677–716.

    Article  Google Scholar 

  129. Smith, A.B. and Jell, P.A., A new cornute carpoid from the Upper Cambrian (Idamean) of Queensland, Mem. Queensl. Mus., 1999, vol. 43, pp. 341–350.

    Google Scholar 

  130. Sumrall, C.D. and Sprinkle, J., Ponticulocarpus, a new cornute-grade stylophoran from the Middle Cambrian Spence Shale of Utah, J. Paleontol., 1999, vol. 73, pp. 886–891.

    Article  Google Scholar 

  131. Sumrall, C.D., Sprinkle, J., and Guensburg, T.E., Systematics and paleocology of Late Cambrian echinoderms from the western United States, J. Paleontol., 1997, vol. 71, pp. 1091–1109.

    Article  Google Scholar 

  132. Sumrall, C.D., Sprinkle, J., Pruss, S., and Finnegan, S., Cardiocystella, a new cornute stylophoran from the Upper Cambrian Whipple Cave Formation, eastern Nevada, USA, J. Paleontol., 2009, vol. 83, pp. 307–312.

    Article  Google Scholar 

  133. Troth, I., The Applications of Palynostratigraphy to the Devonian of Bolivia, PhD Thesis, Southampton, 2006, pp. 1–204 (Unpubl.).

  134. Troth, I., Marshall, J.E.A., Racey, A., and Becker, R.T., Devonian sea-level change in Bolivia: a high palaeolatitude biostratigraphical calibration of the global sea-level curve, Palaeogeogr., Palaeoclimatol., Palaeoecol., 2011, vol. 304, pp. 3–20.

    Article  Google Scholar 

  135. Twenhofel, W.H., Geology of Anticosti Island, Geol. Surv. Can. Mem., 1928, vol. 154, pp. 1–481.

    Google Scholar 

  136. Ubaghs, G., Cothurnocystis Bather, Phyllocystis Thoral and an undetermined member of the order Soluta (Echinodermata, Carpoidea) in the uppermost Cambrian of Nevada, J. Paleontol., 1963, vol. 37, pp. 1133–1142.

    Google Scholar 

  137. Ubaghs, G., Stylophora, in Treatise on Invertebrate Paleontology: Part S. Echinodermata 1(2), Moore, R.C., Ed., Boulder: Geol. Soc. Amer., Boulder, and Lawrence: Univ. Kansas Press, 1967a, pp. S495–S565.

  138. Ubaghs, G., Le genre Ceratocystis Jaekel (Echinodermata, Stylophora), Univ. Kans. Paleontol. Contrib., 1967b, vol. 22, pp. 1–16.

    Google Scholar 

  139. Ubaghs, G., Les Echinodermes “Carpoïdes” de l’Ordovicien Inférieur de la Montagne Noire (France), Paris: Editions du CNRS, 1970, pp. 1–110.

    Google Scholar 

  140. Ubaghs, G., Trois Mitrata (Echinodermata: Stylophora) nouveaux de l’Ordovicien de Tchécoslovaquie, Paläontol. Z., 1979, vol. 53, pp. 98–119.

    Google Scholar 

  141. Ubaghs, G., Réflexions sur la nature et la fonction de l’appendice articulé des “carpoïdes” Stylophora (Echinodermata), Ann. Paléontol., 1981, vol. 67, pp. 33–48.

    Google Scholar 

  142. Ubaghs, G., Deux Stylophora (Homalozoa, Echinodermata) nouveaux pour l’Ordovicien inférieur de la Montagne Noire (France méridionale), Paläontol. Z., 1991, vol. 65, pp. 157–171.

    Book  Google Scholar 

  143. Ubaghs, G., Echinodermes nouveaux (Stylophora, Eocrinoidea) de l’Ordovicien inférieur de la Montagne Noire (France), Ann. Paléontol., 1994, vol. 80, pp. 107–141.

    Google Scholar 

  144. Ubaghs, G., Echinodermes nouveaux du Cambrien supérieur de la Montagne Noire (France méridionale), Geobios, 1998, vol. 31, pp. 809–829.

    Article  Google Scholar 

  145. Ware, D. and Lefebvre, B., Abnormalities in the Lower Ordovician cornute Phyllocystis Thoral (Echinodermata: Stylophora) of Montagne Noire (southern France): implications for plate homology and phylogeny, J. Paleontol., 2007, vol. 81, pp. 779–787.

    Article  Google Scholar 

  146. Woods, I.S. and Jefferies, R.P.S., A new stem-group chordate from the Lower Ordovician of South Wales, and the problem of locomotion in boot-shaped cornutes, Palaeontology, 1992, vol. 35, pp. 1–25.

    Google Scholar 

  147. 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., Vizcaïno, D., and Smith, A.B., Cambrian echinoderm diversity and palaeobiogeography, in Early Palaeozoic Biogeography and Palaeogeography, Harper, D.A.T and Servais, T., Eds, Geol. Soc. London Mem., 2013a, vol. 38, pp. 157–171.

    Google Scholar 

  148. Zamora, S., Zhu, X., and Lefebvre, B., A new Furongian (Cambrian) Echinoderm-Lagerstätte from the Sandu Formation (South China), Cah. Biol. Mar., 2013b, vol. 54, pp. 565–569.

    Google Scholar 

  149. Zhu, X, Peng, S., Zamora, S., Lefebvre, B., and Chen, G., Furongian (upper Cambrian) Guole Konservat-Lagerstätte from South China, Acta Geol. Sin., 2016, vol. 90, pp. 801–808.

    Google Scholar 

Download references

ACKNOWLEDGMENTS

This paper is a contribution to the team “Biosignatures-Vie Primitive” of UMR CNRS 5276 LGLTPE. Field work in Bolivia was supported by the National Geographic Society (grant 6651-99). The authors are particularly grateful to P. Rachebœuf who, during that project, collected all specimens of P. racheboeufi, and provided all necessary information on localities and local stratigraphy. This paper is also a continuation of fieldwork supported by the National Geographic Society (grant 6789-00) and the National Science Foundation (EAR-0205968). The holotype of P.? cybeleae was collected on an Anticosti Island excursion with P. Copper and his students. This study also represents an output of the two European Synthesys projects DE-TAF-4766 and SE-TAF-4765, which made it possible to examine specimens discussed in this paper and deposited, respectively, in the collections of the Museum für Naturkunde, Berlin (Aspidocarpus bohemicus) and the Naturhistoriska Riksmuseet, Stockholm (Lindström’s mitrate). Their respective curators, Christian Neumann and Christina Franzen-Bengtson, are greatly acknowledged for their help and assistance. The authors also thank Thomas E. Guensburg and Sergei V. Rozhnov for their thoughtful reviews. The authors are also particularly grateful to J. Adrien and C. Salaviale for their help with CT-scan investigations of the Anticosti material, N. Podevigne and A. Prieur for photographic assistance, and well as N. Kakavelakis for his help with Ancient Greek. Finally, D. Gendry and E. Robert are thanked for access to important comparison material deposited in the paleontological collections of Rennes 1 University and Lyon 1 University, respectively.

Author information

Authors and Affiliations

  1. UMR CNRS 5276 LGLTPE, bâtiment Géode, campus de la Doua, Université Lyon 1, F-69622, Villeurbanne cedex, France

    B. Lefebvre

  2. School of Earth Sciences, 125 South Oval Mall, The Ohio State University, OH 43210, Columbus, USA

    W. I. Ausich

Authors
  1. B. Lefebvre
    View author publications

    You can also search for this author in PubMed Google Scholar

  2. W. I. Ausich
    View author publications

    You can also search for this author in PubMed Google Scholar

Corresponding authors

Correspondence to B. Lefebvre or W. I. Ausich.

Rights and permissions

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Lefebvre, B., Ausich, W.I. New Siluro-Devonian Anomalocystitids (Echinodermata, Stylophora) from Bolivia and Canada, and a Reevaluation of Skeletal Homologies in Mitrates. Paleontol. J. 55, 932–965 (2021). https://doi.org/10.1134/S0031030121090070

Download citation

  • Received:

  • Revised:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1134/S0031030121090070

Keywords:

Search

Navigation