Current IF 1.5
Latest issue (RSS 2.0)
Contact Editorial Office at
bulletin@geology.cz
Bulletin of Geosciences
Published by ©
Czech Geological Survey,
W. Bohemia Museum Pilsen
Individual sponsors
ISSN: 1802-8225 (online),
1214-1119 (print)

Response of the Silurian jawed polychaete fauna to environmental changes during the Mid-Ludfordian Glaciation in the Prague Basin (Czech Republic)
Published in: Bulletin of Geosciences, volume 100, issue 4; pages: 617 - 644; Received 27 October 2024; Accepted in revised form 19 March 2025; Online 17 May 2025
Keywords: scolecodonts, jawed polychaetes, Silurian, MLCIE, Lau and Kozlowskii bioevents, Prague Basin,
Abstract
Numerous recent studies have revealed that the Ludfordian was an unstable part of the Silurian Period, linked with global climatic changes and glaciation, changes in ocean chemistry, and a distinct perturbation of the carbon cycle expressed by the largest carbon isotope excursion throughout the whole Phanerozoic, the Mid-Ludfordian carbon isotope excursion (MLCIE). These dramatic environmental changes were associated with faunal crises in conodont (Lau conodont Bioevent) and graptolite (Kozlowskii graptolite Bioevent) faunas. The response of other faunal groups to global environmental changes during the mid-Ludfordian has been less thoroughly documented. Here, we present an analysis of the taxonomic composition and stratigraphical distribution of jaw-bearing polychaete worms from the Ludfordian strata of the Prague Basin. Based on more than 5,000 diagnostic posterior maxillae, more than thirty species were recorded from the studied interval. The family Rhytiprionidae was recorded from the peri-Gondwanan region for the first time. During the MLCIE and the Mid-Ludfordian Glaciation interval, the jawed polychaete fauna was disturbed and reorganised but recovered relatively quickly, sooner than, e.g. lingulate brachiopods. The most distinct change was documented at the end of the Polygnathoides siluricus Zone marking the beginning of the glaciation. Approximately half of the species showed a continuous range through the event interval, twenty percent of species represented Lazarus taxa, six percent disappeared and 24 percent appeared during or after the event interval in the Kosov section.References
Andrew, A.S., Hamilton, P.J., Mawson, R., Talent, J.A. & Whitford, D.J. 1994. Isotopic correlation tools in the mid-Palaeozoic and their relation to extinction events. Australian Petroleum Exploration Association (APEA) Journal 34, 268-277.
Barrande, J. 1879. Systeme silurien du centre de la Boheme. Iére partie. Recherches paléontologiques. Vol. 5. Classe des Mollusques. Ordre des Brachiopodes. 226 pp. Privately published, Prague & Paris.
Barrick, J.E, Kleffner, M.A., Gibson, M.A, Peavey, F.N. & Karlsson, H.R. 2010. The mid-Ludfordian Lau Event and Carbon Isotope Excursion (Ludlow, Silurian) in southern Laurentia - Preliminary Results. Bollettino della Societa Paleontologica Italiana 49(1), 13-33.
Beesley, P.L., Ross, G.J.B. & Glasby, C.J. 2000. Fauna of Australia - Volume 4A Polychaetes & Allies. 465 pp. Australian Biological Resources Study/CSIRO Publishing.
Bergman, C. 1979. Polychaete jaws. Lower Wenlock faunal and floral dynamics - Vattenfallet section, Gotland. Sveriges Geologiska Undersökning 73(3), 92-102.
Bergman, C.F. 1987. Silurian paulinitid jawed polychaetes from Gotland. 145 pp. Ph.D. thesis, University of Lund, Sweden.
Bergman, C.F. 1989. Silurian paulinitid polychaetes from Gotland. Fossils and Strata 25, 1-128.
Bergman, C.F. 1995. Symmetroprion spatiosus (Hinde), a jawed polychaete showing preference for reef environments in the Silurian of Gotland. GFF 117(3), 143-150.
Bergman, C.F., Eriksson, M.E. & Calner, M. 2003. The jawed annelid Rhytiprion magnus Kielan-Jaworowska, 1966 - a muddy bottom dweller. GFF 125(4), 191-199.
Berner, R.A. 2005. The rise of trees and how they changed Paleozoic atmospheric CO2, climate and geology, 1-7. In Baldwin, I.T., Caldwell, M.M., Heldmaier, G., Jackson, R.B., Lange, O.L., Mooney, H.A., Schulze, E.-D., Sommer, U., Ehleringer, J.R., Denise Dearing, M. & Cerling, T.E. (eds) A History of Atmospheric CO2 and Its Effects on Plants, Animals, and Ecosystems, Ecological Studies 177. Springer.
Berner, R.A. 2006. GEOCARBSULF: A combined model for Phanerozoic atmospheric O2 and CO2. Geochimica et Cosmochimica Acta 70(23), 5653-5664.
Berner, R.A. 2009. Phanerozoic atmospheric oxygen: New results using the GEOCARBSULF model. American Journal of Science 309(7), 603-606.
Berner, R.A., Beerling, D.J., Dudley, R., Robinson, J.M. & Wildman, R.A. 2003. Phanerozoic atmospheric oxygen. Annual Review of Earth and Planetary Sciences 31(1), 105-134.
Bickert, T., Pätzold, J., Samtleben, C. & Munnecke, A. 1997. Paleoenvironmental changes in the Silurian indicated by stable isotopes in brachiopod shells from Gotland, Sweden. Geochimica et Cosmochimica Acta 61, 2717-2730.
Bowman, C.N., Young, S.A., Kaljo, D., Eriksson, M.E., Them II, T.R., Hints, O., Martma, T. & Owens, J. 2019. Linking the progressive expansion of reducing conditions to a stepwise mass extinction event in the late Silurian oceans. Geology 47, 968-972.
Bowman, C.N., Lindskog, A., Kozik, N.P., Richbourg, C.G., Owens, J.D. & Young, S.A. 2020. Integrated sedimentary, biotic, and paleoredox dynamics from multiple localities in southern Laurentia during the late Silurian (Ludfordian) extinction event. Palaeogeography, Palaeoclimatology, Palaeoecology 553, 109799.
Bowman, C.N., Them II, T.R., Knight, M.D., Kaljo, D., Eriksson, M.E., Hints, O., Martma, T., Owens, J.D. & Young, S.A. 2021. A multi-proxy approach to constrain reducing conditions in the Baltic Basin during the late Silurian Lau carbon isotope excursion. Palaeogeography, Palaeoclimatology, Palaeoecology 581, 110624.
Branson, E.B. & Mehl, M.G. 1933. Conodont studies. University of Missouri Studies 8(1-4), 1-343.
Brett, C.E. & Walker, S.E. 2002. Predators and predation in Paleozoic marine environments. Paleontological Society Papers 8, 93-118.
Brett, C.E., Ferreti, A., Histon, K. & Schönlaub, H.P. 2009. Silurian sequence stratigraphy of the Carnic Alps, Austria. Palaeogeography, Palaeoclimatology, Palaeoecology 279, 1-28.
Chlupáč, I., Havlíček, V., Kříž, J., Kukal, Z. & Štorch, P. 1998. Palaeozoic of the Barrandian (Cambrian to Devonian). 183 pp. Czech Geological Survey, Prague.
Clack, J.A. 2007. Devonian climate change, breathing, and the origin of the tetrapod stem group. Integrative and Comparative Biology 47, 510-523.
Cocks, L.R.M. & Torsvik, T.H. 2002. Earth geography from 500 to 400 million years ago: a faunal and palaeomagnetic review. Journal of the Geological Society of London 159, 631-644.
Cocks, L.R.M. & Torsvik, T.H. 2013. New global palaeogeographical reconstructions for the Early Palaeozoic and their generation, 5-24. In Harper, D.A.T. & Servais, T. (eds) Early Palaeozoic Biogeography and Palaeogeography. Geological Society London, Memoir 38.
Conway Morris, S. 1979. Middle Cambrian polychaetes from the Burgess Shale of British Columbia. Philosophical Transactions of the Royal Society London B285, 227-274.
Courtinat, B. 1998. New genera and new species of scolecodonts (fossil annelids) with paleoenvironmental and evolutionary considerations. Micropaleontology 44(4), 435-440.
Cramer, B.D. & Saltzman, M.R. 2007. Fluctuations in epeiric sea carbonate production during Silurian positive carbon isotope excursions: A review of proposed paleoceanographic models. Palaeogeography, Palaeoclimatology, Palaeoecology 245, 37-45.
Cramer, B.D., Brett, C.E., Melchin, M.J., Männik, P., Kleffner, M.A., McLaughlin, P.I., Lloydell, D.K., Munnecke, A., Jeppsson, L., Corradini, C., Brunton, F.R. & Saltzman, M.R. 2011. Revised correlation of Silurian provincial series of North America with global and regional chronostratigraphic and δ13Ccarb chemostratigraphy. Lethaia 44, 185-202.
Díaz-Castaneda, V. & Reish, D.J. 2009. Chapter 11 Polychaetes in Environmental Studies, 205-227. In Shain, D.H. (ed.) Annelids in Modern Biology. Wiley-Blackwell.
Eisenack, A. 1975. Beiträge zur Anneliden-Forschung, I. Neues Jahrbuch für Geologie und Paläontologie. Abhandlungen 150, 227-252.
Eriksson, M. 1997. Lower Silurian polychaetaspid polychaetes from Gotland, Sweden. GFF 119, 213-230.
Eriksson, M.E. 1998. Dubichaetaspis bergmani gen. et sp. n., a facies controlled polychaete from the Silurian of Gotland, Sweden. GFF 120(1), 21-25.
Eriksson, M. 2001. Silurian Ramphoprionid polychaetes from Gotland, Sweden. Journal of Paleontology 75(5), 993-1015.
Eriksson, M.E. 2006a. Polychaete jaw apparatuses and scolecodonts from the Silurian Ireviken Event interval of Gotland, Sweden. GFF 128(2), 97-101.
Eriksson, M.E. 2006b. The Silurian Ireviken Event and vagile benthic faunal turnovers (Polychaeta; Eunicida) on Gotland, Sweden. GFF 128(2), 91-95.
Eriksson, M.J. & Calner, M. 2008. A sequence stratigraphical model for the late Ludfordian (Silurian) of Gotland, Sweden - implications for timing between changes in sea-level, palaeoecology, and the global carbon cycle. Facies 54, 253-276.
Eriksson, M.E. & Frisk, Å.M. 2011. Polychaete palaeoecology in an early Late Ordovician marine astrobleme of Sweden. Geological Magazine 148(2), 269-287.
Eriksson, M.E. & Hints, O. 2009. Vagrant benthos (Annelida; Polychaeta) associated with Upper Ordovician carbonate mud-mounds of subsurface Gotland, Sweden. Geological Magazine 146(3), 451-462.
Eriksson, M.E., Bergman, C.F. & Jeppsson, L. 2004. Silurian scolecodonts. Review of Palaeobotany and Palynology 131, 269-300.
Eriksson, M.E., Nilsson, E.K. & Jeppsson, L. 2009. Vertebrate extinctions and reorganizations during the Late Silurian. Geology 37(8), 739-742.
Eriksson, M.E., Grahn, Y., Bosetti, E.P. & Vega, C.S. 2011. Malvinokaffric Realm polychaetes from the Devonian Ponta Grossa Formation, Parana Basin (southern Brazil), with a discussion and re-evaluation of the species described by Lange, 117-150. In Bosetti, E.P., Grahn, Y. & Melo, J.H.G. (eds) Essays in honour of Frederico Waldemar Lange, pioneer of Brazilian micropaleontology. Interciencia.
Eriksson, M.E., Hints, O., Paxton, H. & Tonarová, P. 2013. Ordovician and Silurian polychaete diversity and biogeography, 265-272. In Harper, D.A.T., Servais, T. (eds) Early Palaeozoic Biogeography and Palaeogeography. Geological Society London, Memoir 38.
Farkaš, J., Frýda, J. & Holmden, C. 2016. Calcium isotope constraints on the marine carbon cycle and CaCO3 deposition during the late Silurian (Ludfordian) positive δ13C excursion. Earth and Planetary Science Letters 451, 31-40.
Farkaš. J., Wallmann, K., Mosley, L., Staudigel, P., Zheng, X.-Y., Leyden, E., Shao, Y., Frýda, J., Holmden, C. & Eisenhauer, A. 2024. Alkalinity and elemental cycles in present and past ocean: Insight from Geochemical Modeling and Alkali and Alkaline Earth Metal Isotopes, 33-87. In Reference Module in Earth Systems and Environmental Sciences 2024.
Fatka, O.& Mergl, M. 2009. The “microcontinent” Perunica: status and story 15 years after conception. Geological Society London, Special Publication 325(1), 65-101.
Franke, W., Cocks, L.R.M. & Torsvik, T.H. 2017. The Palaeozoic Variscan oceans revisited. Gondwana Research 48, 257-284.
Frýda, J. & Manda, Š. 2013. A long-lasting steady period of isotopically heavy carbon in the late Silurian ocean: evolution of the δ13C record and its significance for an integrated δ13C, graptolite and conodont stratigraphy. Bulletin of Geosciences 88(2), 463-482.
Frýda, J., Lehnert, O., Frýdová, B., Farkaš, J. & Kubajko, M. 2021a. Carbon and sulfur cycling during the mid-Ludfordian anomaly and the linkage with the late Silurian Lau/Kozlowskii Bioevent. Palaeogeography, Palaeoclimatology, Palaeoecology 564, 110152.
Frýda, J., Lehnert, O., Joachimski, M.M., Männik, P., Kubajko, M., Mergl, M., Farkaš, J. & Frýdová, B. 2021b. The Mid-Ludfordian (late Silurian) Glaciation: a link with global changes in ocean chemistry and ecosystem overturns. Earth-Science Reviews 220, 103652.
Giangrande, A., Licciano, M. & Musco, L. 2005. Polychaetes as environmental indicators revisited. Marine Pollution Bulletin 50, 1153-1162.
Gocke, M., Lehnert, O. & Frýda, J. 2012. Facies development across the Late Silurian Lau Event based on temperate carbonates of the Prague Basin (Czech Republic). Facies 59, 611-630.
Hartke, E.R., Cramer, B.D., Calner, M., Melchin, M.J., Barnett, B.A., Oborny, S.C. & Bancroft, A.M. 2021. Decoupling δ13Ccarb and δ13Corg at the onset of the Ireviken Carbon Isotope Excursion: Δ13C and organic carbon burial (forg) during a Silurian oceanic anoxic event. Global and Planetary Change 196, 103373.
Havlíček, V. & Štorch, P. 1990. Silurian brachiopods and benthic communities in the Prague Basin (Czechoslovakia). Rozpravy Ústředního ústavu geologického 48, 1-275.
Havlíček, V. & Štorch, P. 1999. Silurian and Lochkovian communities of the Prague Basin (Barrandian area, Czechoslovakia), 200-228. In Boucot, A.J. & Lawson, J.D. (eds) Final report, Project Ecostratigraphy. Paleocommunities: A case study from the Silurian and Lower Devonian. Cambridge University Press, Cambridge.
Havlíček,V., Vaněk, J. & Fatka, O. 1994. Perunica microcontinent in the Ordovician (its position within the Mediterranean Province, series division, benthic and pelagic associations). Sborník geologických věd, Paleontologie 46, 23-56.
Hayes, J.M., Strauss, H. & Kaufman, A.J. 1999. The abundance of 13C in marine organic matter and isotopic fractionation in the global biogeochemical cycle of carbon during the past 800 Ma. Chemical Geology 161, 103-125.
Hinde, G.J. 1879. On Annelid jaws from the Cambro-Silurian, Silurian and Devonian formations in Canada and from Lower Carboniferous in Scotland. The Quarterly Journal of the Geological Society of London 35, 370-389.
Hinde, G.J. 1880. On Annelid jaws from the Wenlock and Ludlow formations of the West of England. Quarterly Journal of the Geological Society of London 36, 368-378.
Hinde, G.J. 1882. On Annelid remains from the Silurian strata of the Isle of Gotland. Bihang till Kongliga Vetenskaps-Akademiens Handlingar, Bihang till Kungliga Vetenskapsakademiens Handlingar 7(5), 1-28.
Hints, L., Hints, O., Kaljo, D., Kiipli, T., Männik, P., Nolvak, J. & Pärnaste, H. 2010. Hirnantian (latest Ordovician) bio- and chemostratigraphy of the Stirnas-18 core, western Latvia. Estonian Journal of Earth Sciences 59(1), 1-24.
Hints, O. 1998a. Revision of the Ordovician and Silurian ramphoprionid polychaetes from Severnaya Zemlya, Russian Arctic. Proceedings of the Estonian Academy of Sciences, Geology 47(2), 77-85.
Hints, O. 1998b. Late Viruan (Ordovician) polychaete jaws from North Estonia and the St. Petersburg Region. Acta Palaeontologica Polonica 43(3), 471-516.
Hints, O. 1999. Two new polychaete families from the Upper Ordovician of Estonia. Palaeontology 42(5), 897-906.
Hints, O. 2000. Ordovician eunicid polychaetes of Estonia and surrounding areas: review of their distribution and diversification. Review of Palaeobotany and Palynology 113(1-3), 41-55.
Hints, O. 2001. Distribution of scolecodonts. Valga (10) drill core. Estonian Geological Sections Bulletin 3, 12-14.
Hints, O. & Erikssson, M.E. 2007a. Diversification and biogeography of scolecodont-bearing polychaetes in the Ordovician. Palaeogeography, Palaeoclimatology, Palaeoecology 245(1-2), 95-114.
Hints, O. & Erikssson, M.E. 2007b. Biogeography of Ordovician and Silurian jaw-bearing polychaetes. Acta Palaeontologica Sinica, Supplement 46, 181-187.
Hints, O. & Eriksson, M. 2010. Ordovician polychaeturid polychaetes: Taxonomy, distribution and palaeoecology. Acta Palaeontologica Polonica 55(2), 309-320.
Hints, O. & Tonarová, P. 2023. A diverse Hirnantian scolecodont assemblage from northern Estonia and resilience of polychaetes to the end-Ordovician mass extinction. Estonian Journal of Earth Sciences 72(1), 46-49.
Hints, O., Bergman, C.F.& Märss, T. 2000. Silurian jawed polychaetes from Cornwallis and Baillie-Hamilton islands, Canadian Arctic. Pan-Arctic Palaeozoic Tectonics, Evolution of Basins and Faunas. Ichthyolith Issues, Special Publication 6, 35-38.
Hints, O., Hints, L., Meidla, T. & Sohar, K. 2003. Biotic effects of the Ordovician Kinnekulle ashfall recorded in northern Estonia. Bulletin of Geological Society of Denmark 50, 115-123.
Hints, O., Eriksson, M., Högström, A.E.S., Kraft, P. & Lehnert, O. 2004. Chapter 23. Worms, Worm-like and Sclerite-bearing Taxa, 223-230. In Webby, B.D., Paris, F., Droser, M.L. & Percival, I.G. (eds) The Great Ordovician Biodiversification Event. Columbia University Press.
Hints, O., Killing, M., Männik, P. & Nestor, V. 2006. Frequency patterns of chitinozoans, scolecodonts, and conodonts in the upper Llandovery and lower Wenlock of the Paatsalu core, western Estonia. Proceedings of the Estonian Academy of Sciences 55(2), 128-155.
Hints, O., Tonarová, P. & Desrochers, A. 2016. Late Ordovician jaw-bearing polychaetes from Anticosti Island, eastern Canada, and their biogeographic significance. Canadian Journal of Earth Sciences 53(7), 731-738.
Hints, O., Tonarová, P., Eriksson, M.E., Rubinstein, C.V. & Puente, G.S. de la 2017. Early Middle Ordovician scolecodonts from north-western Argentina and the emergence of labidognath polychaete jaw apparatuses. Palaeontology 60(4), 583-593.
Histon, K. & Schönlaub, H.P. 1999. The Palaeozoic of the Southern Alps. Berichte der Geologisches Bundesanstalt 47, 6-30.
Holmden, C., Creaser, R.A., Muehlenbachs, K., Leslie, S.A. & Bergström, S.M. 1998. Isotopic evidence for geochemical decoupling between ancient epeiric seas and bordering oceans: Implications for secular curves. Geology 26, 567-570.
Holmden, C., Panchuk, K. & Finney S.C. 2012. Tightly coupled records of Ca and C isotope changes during the Hirnantian glaciation in an epeiric sea setting. Geochimica et Cosmochimica Acta 98, 94-106.
Horný, R. 1955a. Studie o vrstvách budňanských v západní části barrandienského siluru. Sborník Ústředního ústavu geologického 21, 315-448.
Horný, R., 1955b. Báse vrstev kopaninských eβ1 na jihozápadním okraji vulkanické facie (Kosov u Berouna). Věstník Ústředního ústavu geologického 30, 81-86.
Hutchings, P. 1998. Biodiversity and functioning of polychaetes in benthic sediments. Biodiversity and Conservation 7, 1133-1145.
Hutchings, P.A. & Fauchald, K. 2000. Class Polychaeta. Definition and General Description, 4-7. In Beesley, P.L., Ross, G.J.B. & Glasby, C.J., (eds) Polychaetes & Allies: The Southern Synthesis. Fauna of Australia. CSIRO Publishing, Melbourne.
Ireland, H.A. 1961. New phosphatic brachiopods from the Silurian of Oklahoma. Journal of Paleontology 35, 1137-1142.
Jarochowska, E., Nohl, T., Grohganz, M., Hohmann, N., Vandenbroucke, T.R.A. & Munnecke, A. 2020. Reconstructing depositional rates and their effect on paleoenvironmental proxies: The case of the Lau Carbon Isotope Excursion in Gotland, Sweden. Paleoceanography and Paleoclimatology 35, e2020PA003979.
Jeppsson, L. 1987. Lithological and conodont distributional evidence for episodes of anomalous oceanic conditions during the Silurian, 129-145. In Aldridge, R.J. (ed.) Palaeobiology of Conodonts. Ellis Horwood Ltd, Chichester, England.
Jeppsson, L. 1990. An oceanic model for lithological and faunal changes. Journal of the Geological Society London 147, 663-674.
Jeppsson, L. 1993. Silurian events, the theory, and the conodonts. Proceedings of the Estonian Academy of Sciences Geology 42, 23-27.
Jeppsson, L. & Aldridge, R.J. 2000. Ludlow (late Silurian) oceanic episodes and events. Journal of the Geological Society London 157, 1137-1148.
Jeppsson, L., Talent, J.A., Mawson, R., Simpson, A.J., Andrew, A., Calner, M., Whitford, D., Trotter, J.A., Sandström, O. & Caldon, H.J. 2007. High-resolution Late Silurian correlations between Gotland, Sweden, and the Broken River region, NE Australia: lithologies, conodonts and isotopes. Palaeogeography, Palaeoclimatology, Palaeoecology 245, 115-137.
Jeppsson, L., Talent, J.A., Mawson, R., Andrew, A., Corradini, C., Simpson, A.J., Wigforss-Lange, J. & Schönlaub, H.P. 2012. Late Ludfordian correlations and the Lau Event, 653-675. In Talent, J.A. (ed.) Earth and Life: Global Biodiversity, Extinction Intervals and Biogeographic Perturbations Through Time. Springer Netherlands.
Kaljo, D., Boucot, A.J., Corfield, R.M., Le Herisse, A., Koren, T.N., Kříž, J., Männik, P., Märss, T., Nestor, V., Shaver, R.H., Siveter, D.J. & Viira, V. 1995. Silurian bioevents, 173-224. In Walliser, O.H. (ed.) Global events and event stratigraphy in the Phanerozoic. Springer-Verlag.
Kaljo, D., Boucot, A.J., Corfield, R.M., Herisse, A.L., Koren’, T.N., Kriz, J., Männik, P., Märss, T., Nestor, V., Shaver, R.H., Siveter, D.J. & Viira, V. 1996. Silurian Bio-Events, 173-224. In Walliser, O.H. (ed.) Global Events and Event Stratigraphy in the Phanerozoic. Springer Berlin Heidelberg.
Kaljo, D., Kiipli, T. & Martma, T. 1997. Carbon isotope event markers through the Wenlock-Pridoli sequence at Ohesaare (Estonia) and Priekule (Latvia). Palaeogeography, Palaeoclimatology, Palaeoecology 132(1-4), 211-223.
Kaljo, D., Grytsenko, V., Martma, T. & Motus, M.A. 2007. Three global carbon isotope shifts in the Silurian of Podolia (Ukraine): stratigraphical implications. Estonian Journal of Earth Sciences 56, 205-220.
Kaljo, D., Brazauskas, A., Kaminskas, D. & Martma, T. 2011. New developments in Baltic Silurian chemostratigraphy: results and some tasks, 31. In Lukshevics, E., Stinkulis, G. & Vasilkova, J. (eds) The Eight Baltic Stratigraphical Conference. Abstracts. University of Latvia.
Kaljo, D., Grytsenko, V., Kallaste, T., Kiipli, T. & Martma, T. 2014. Upper Silurian stratigraphy of Podolia revisited: carbon isotopes, bentonites and biostratigraphy. GFF 136, 136-141.
Kaljo, D., Einasto, R., Martma, T., Märss, T., Nestor, V. & Viira, V. 2015. A bio-chemostratigraphical test of the synchroneity of biozones in the upper Silurian of Estonia and Latvia with some implications for practical stratigraphy. Estonian Journal of Earth Sciences 64, 267-283.
Kielan-Jaworowska, Z. 1961. On two Ordovician polychaete jaw apparatuses. Acta Palaeontologica Polonica 6(3), 237-254.
Kielan-Jaworowska, Z. 1962. New Ordovician genera of polychaete jaw apparatuses. Acta Palaeontologica Polonica 7(3-4), 291-325.
Kielan-Jaworowska, Z. 1966. Polychaete jaw apparatuses from the Ordovician and Silurian of Poland and a comparison with modern forms. Palaeontologia Polonica 16, 1-152.
Klug, C., Kröger, B., Kiessling, W., Mullins, G.L., Servais, T., Frýda, J., Korn, D. & Turner, S. 2010. The Devonian nekton revolution. Lethaia 43, 465-477.
Kozłowski, R. 1956. Sur quelques appareils masticateurs des Annelides Polychetes ordoviciens. Acta Palaeontologica polonica 1(3), 165-210.
Kozłowski, W. 2015. Eolian dust influx and massive whitings during the Kozlowski/Lau Event: carbonate hypersaturation as a possible driver of the mid-Ludfordian carbon isotope excursion. Bulletin of Geosciences 90, 807-840.
Kozłowski, W. 2020. Marine snow and epipelagic suspensoids in the Reda carbonates and a pronounced mid-Ludfordian (Silurian) CIE in the axis of the Baltic Basin (Poland). Acta Geologica Polonica 70(4), 529-567.
Kozłowski, W. & Munnecke, A. 2010. Stable carbon isotope development and sea-level changes during the Late Ludlow (Silurian) of the Łysogóry region (Rzepin section, Holy Cross Mountains, Poland). Facies 56, 615-633.
Kozłowski, W. & Sobień, K. 2012. Mid-Ludfordian coeval carbon isotope, natural gamma ray and magnetic susceptibility excursions in the Mielnik IG-1 borehole (Eastern Poland)-Dustiness as a possible link between global climate and the Silurian carbon isotope record. Palaeogeography, Palaeoclimatology, Palaeoecology 339-341, 74-97.
Kříž, J. 1991. The Silurian of the Prague Basin (Bohemia): Tectonic, eustatic, and volcanic controls on facies and faunal development. Special Papers in Palaeontology 44, 179-204.
Kříž, J. 1992. Silurian Field Excursions: Prague Basin (Barrandian), Bohemia. 111 pp. National Museum of Wales, Cardiff, Geological Series 13.
Kříž, J. 1998a. Silurian, 70-101. In Chlupáč, I., Havlíček, V., Kříž, J., Kukal, Z. & Štorch, P. (eds) Palaeozoic of the Barrandian. Czech Geological Survey, Prague.
Kříž, J. 1998b. Recurrent Silurian-Lowest Devonian cephalopod limestones of Gondwanan Europe and Perunica, 183-198. In Landing, E. & Johnson, M.E. (eds) Silurian Cycles: Linkages of Dynamic Stratigraphy with Atmosphaeric, Oceanic, and Tectonic Changes (James Hall Centennial Volume). New York State Museum Bulletin 491.
Kump, L. & Arthur, M.A. 1999. Interpreting carbon-isotope excursions: carbonates and organic matter. Chemical Geology 161, 181-198.
Kump, L.R., Arthur, M.A., Patzkowsky, M.E., Gibbs, M.T., Pinkus, M.T. & Sheehan, P.M. 1999. A weathering hypothesis for glaciation at high atmospheric pCO2 during the Late Ordovician. Palaeogeography, Palaeoclimatology, Palaeoecology 152(1-2), 173-187.
Lange, F.W. 1947. Annelidos poliquetas dos folhelhos devonianos do Paraná. Arquivos do Museu Paranaese 6(5), 161-230.
LaPorte, D.F., Holmden, C., Patterson, W.P., Loxton, J.D., Melchin, M.J., Mitchell, C., Finney, S.C. & Sheets, H.D. 2009. Local and global perspectives on carbon and nitrogen cycling during the Hirnantian Glaciation. Palaeogeography, Palaeoclimatology, Palaeoecology 276, 182-195.
Lehnert, O., Frýda, J., Buggisch, W. & Manda, Š. 2003. A first report of the Ludlow Lau event from the Prague Basin (Barrandian, Czech Republic), 139-144. In Albanesi, G.L., Beresi, M.S. & Peralta, S.H. (eds) INSUGEO, Serie Correlación Geológica 18.
Lehnert, O., Eriksson, M.J., Calner, M., Joachimski, M. & Buggisch, W. 2007a. Concurrent sedimentary and isotopic indications for global climatic cooling in the Late Silurian. Acta Palaeontologica Sinica 46, 249-255.
Lehnert, O., Frýda, J., Buggisch, W., Munnecke, A., Nützel, A., Kříž, J. & Manda, Š. 2007b. δ13C records across the late Silurian Lau event: New data from middle palaeolatitudes of northern peri-Gondwana (Prague Basin, Czech Republic) Palaeogeography, Palaeoclimatology, Palaeoecology 245, 227-244.
Lehnert, O., Männik, P., Joachimski, M.M., Calner, M. & Frýda, J. 2010. Palaeoclimate perturbations before the Sheinwoodian glaciation: A trigger for extinctions during the ’Ireviken Event’. Palaeogeography, Palaeoclimatology, Palaeoecology 296(3-4), 320-331.
Lin, C.Y., Turchyn, A., Steiner, Z., Bots, P., Lampronti, G.I. & Tosca, N.J. 2018. The role of microbial sulfate reduction in calcium carbonate polymorph selection. Geochimica et Cosmochimica Acta 237, 184-204.
Loydell, D.K. 2007. Early Silurian positive δ13C excursions and their relationship to glaciations, sea-level changes and extinction events. Geological Journal 42, 531-546.
Loydell, D.K. & Frýda, J. 2011. At what stratigraphical level is the mid Ludfordian (Ludlow, Silurian) positive carbon isotope excursion in the type Ludlow area, Shropshire, England? Bulletin of Geosciences 86, 197-208.
Manda, Š. & Frýda, J. 2014. Evolution of the late Ludlow to early Lochkovian brachiopod, trilobite and bivalve communities of the Prague Basin and their link with the global carbon cycle. GFF 136(1), 179-184.
Manda, Š. & Kříž, J. 2006. Environmental and biotic changes in subtropical isolated carbonate platforms during the late Silurian Kozlowskii Event, Prague Basin. GFF 128(2), 161-168.
Manda, Š., Štorch, P., Slavík, L., Frýda, J., Kříž, J. & Tasáryová, Z. 2012. The graptolite, conodont and sedimentary record through the late Ludlow Kozlowskii Event (Silurian) in the shale-dominated succession of Bohemia. Geological Magazine 149(3), 507-531.
Männik, R., Ainsaar, L., Männik, P., Meidla, T., Radzevičius, S. & Hints, O. 2024. Microfossil response to the late Silurian Lau Event in the Bebirva-111 drill core, Lithuania, 26. In Hints, O., Männik, P. & Toom, U. (eds) XI Baltic Stratigraphical Conference. Abstracts and Field Guide. Geological Society of Estonia.
Männil, R.M. & Zaslavskaya, N.M. 1985. Silurian polychaetes from northern Siberia. 98-119, 127-130. In Gudina, V.I. & Kanigin, A.V. (eds) Phanerozoic microfauna from Siberia with surroundings. Trudy Instituta Geologii I Geofiziki 615.
Märss, T. 1992. Vertebrate history in the Late Silurian. Proceedings of the Estonian Academy of Sciences, Geology 41, 205-214.
McKerrow, W.S., Mac Niocaill, C., Ahlberg, P., Clayton, G., Cleal, C.J. & Eagar, M.C. 2000. The Late Palaeozoic relations between Gondwana and Laurasia. Geological Society London Special Publications 179(1), 9-20.
Melchin, M.J. & Holmden, C. 2006. Carbon isotope chemostratigraphy in Arctic Canada. Sea-level forcing of carbonate platform weathering and implications for Hirnantian global correlation. Palaeogeography, Palaeoclimatology, Palaeoecology 234, 186-200.
Melchin, M.J., Sadler, P.M. & Cramer, B.D. 2020. Chapter 21 - The Silurian period, 695-732. In Gradstein, F.M., Ogg, J.G., Schmitz, M.D. & Ogg, G.M. (eds) The Geological Time Scale 2020. Elsevier B.V.
Mergl, M. 1982. Caenotreta (Inarticulata, Brachiopoda) in the Upper Silurian of Bohemia. Věstník Ústředního ústavu geologického 57, 115-116.
Mergl, M., Frýda, J. & Kubajko, M. 2018. Response of organophosphatic brachiopods to the mid-Ludfordian (late Silurian) carbon isotope excursion and associated extinction events in the Prague Basin (Czech Republic). Bulletin of Geosciences 93(3), 369-400.
Mierzejewski, P. 1978. New placognath Eunicida (Polychaeta) from the Ordovician and Silurian of Poland. Acta Geologica Polonica 28(2), 273-281.
Munnecke, A., Samtleben, C. & Bickert, T. 2003. The Ireviken Event in the lower Silurian of Gotland, Sweden - relation to similar Palaeozoic and Proterozoic events. Palaeogeography, Palaeoclimatology, Palaeoecology 195(1-2), 99-124.
Nützel, A. & Frýda, J. 2003. Palaeozoic plankton revolution: Evidence from early gastropod ontogeny. Geology 31(9), 829-831.
Olsgard, F., Brattegard, T. & Holthe, T. 2003. Polychaetes as surrogates for marine biodiversity: lower taxonomic resolution and indicator groups. Biodiversity and Conservation 12, 1033-1049.
Panchuk, K.M., Holmden, C. & Leslie, S.A. 2006. Local controls on carbon cycling in the Ordovician Midcontinent region of North America with implications for carbon isotope secular curves. Journal of Sedimentary Research 76, 200-211.
Paxton, H. 2000. Family Eunicidae, 91-94. In Beesley, P.L., Ross, G.J.B. & Glasby, C.J. (eds) Polychaetes & Allies: The Southern Synthesis. Fauna of Australia Vol. 4A Polychaeta, Myzostomida, Pogonophora, Echiura, Sipuncula. Melbourne, CSIRO Publishing.
Paxton, H. 2009. Phylogeny of Eunicida (Annelida) based on morphology of jaws. Zoosymposia 2, 241-264.
Pisarzowska, A., Becker, R.T., Aboussalam, Z.S., Szczerba, M., Sobień, K., Kremer, B., Owocki, K. & Racki, G. 2020. Middlesex/punctata Event in the Rhenish Basin (Padberg section, Sauerland, Germany) - Geochemical clues to the early-middle Frasnian perturbation of global carbon cycle. Global and Planetary Change 191, 103211.
Pšenička, J., Bek, J., Frýda, J., Žárský, V., Uhlířová, M. & Štorch, P. 2021. Dynamics of Silurian plants as response to climate changes. Life 11(9), 906.
Rey, A. del, Havsteen, J.C., Bizzarro, M. & Dahl, T.W. 2020. Untangling the diagenetic history of uranium isotopes in marine carbonates: a case study tracing the δ238U composition of late Silurian oceans using calcitic brachiopod shells. Geochimica et Cosmochimica Acta 287, 93-110.
Rey, Á. del, Frýda, J., Calner, M., Frýdová, B., Zhang, F., Wang, Ch., Planavsky, N. & Dahl, T.W. 2023. Mid-Ludfordian uranium isotope records distinguish periods of expansive marine anoxia and carbonate hypersaturation during the late Silurian Lau/Kozlowskii bioevent. Planetary and Global Changes 229(1-9), 104248.
Saltzman, M.R. & Thomas, E. 2012. Carbon isotope stratigraphy, 207-232. In Gradstein, F., Ogg, J., Schmitz, M.D. & Ogg, G. (eds) The Geologic Time Scale 2012. Elsevier.
Samtleben, C., Munnecke, A., Bickert, T. & Pätzold, J. 1996. The Silurian of Gotland (Sweden): Facies interpretation based on stable isotopes in brachiopod shells. Geologische Rundschau 85, 278-292.
Schachat, S.R., Labandeira, C.C., Saltzman, M.R., Cramer, B.D., Payne, J.L. & Boyce, C.K. 2018. Phanerozoic pO2 and the early evolution of terrestrial animals. Proceedings of the Royal Society B 285(1871), 20172631.
Schrag, P.D., Higgins, J.A., Macdonald, F.A. & Johnston, D. 2013. Authigenic Carbonate and the History of Global Carbon Cycle. Science 339, 540-543.
Scotese, C.R. 2014. Atlas of Silurian and Middle-Late Ordovician Paleogeographic Maps (Mollweide Projection), Maps 73-80, Volume 5. The Early Paleozoic, PALEOMAP Atlas for ArcGIS, PALEOMAP Project, Evanston, IL.
Scott, D.M. 1976. Circadian rhythms of anaerobiosis in a polychaete annelid. Nature 262, 811-813.
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 London, Special Publication 325(1).
Sheehan, P.M. 2001. The Late Ordovician Mass Extinction. Annual Reviews of Earth and Planetary Sciences 29, 331-364.
Signor, P.W. & Brett, C.E. 1984. The mid-Paleozoic precursor to the Mesozoic marine revolution. Paleobiology 10, 229-245.
Slavík, L. & Carls, P. 2012. Post-Lau Event (late Ludfordian, Silurian) recovery of conodont faunas of Bohemia. Bulletin of Geosciences 87(4), 815-832.
Slavík, L., Kříž, J. & Carls, P. 2010. Reflection of the mid-Ludfordian Lau Event in conodont faunas of Bohemia. Bulletin of Geosciences 85(3), 395-414.
Šnajdr, M. 1951. On Errant Polychaeta from the Lower Paleozoic of Bohemia. Sborník Ústředního ústavu geologického, Paleontology 18, 241-296.
Spiridonov, A., Stankevič, R., Gečas, T., Šilinskas, T., Brazauskas, A., Meidla, T., Ainsaar, L., Musteikis, P. & Radzevičius, S. 2017. Integrated record of Ludlow (Upper Silurian) oceanic geobioevents - Coordination of changes in conodont, and brachiopod faunas, and stable isotopes. Gondwana Research 51, 272-288.
Spiridonov, A., Samsone, J., Brazauskas, A., Stankevič, R., Meidla, T., Ainsaar, L. & Radzevičius, S. 2020. Quantifying the community turnover of the uppermost Wenlock and Ludlow (Silurian) conodonts in the Baltic Basin. Palaeogeography, Palaeoclimatology, Palaeoecology 549, 109128.
Sproson, A.D., Pogge von Strandmann, P.A.E., Selby, D., Jarochowska, E., Frýda, J., Hladil, J., Loydell, D.K., Slavík, L., Calner, M., Maier, G., Munnecke, A. & Lenton, T.M. 2022. Osmium and lithium isotope evidence for weathering feedbacks linked to orbitally paced organic carbon burial and Silurian glaciations. Earth and Planetary Science Letters 577, 117260.
Stankevič, R., Venckute-Aleksiene, A., Radzevičius, S. & Spiridonov, A. 2024. Phytoplankton and zooplankton paleocommunity change before and during the onset of the Lau Extinction Event (Ludlow, Silurian). Marine Micropaleontology 189, 102368.
Stauffer, C.R. 1933. Middle Ordovician Polychaeta from Minnesota. Bulletin of the Geological Society of America 44, 1173-1218.
Steemans, P., Petus, E., Breuer, P., Mauller-Mendlowicz, P. & Gerienne, P. 2012. Palaeozoic innovations in the micro- and megafossil plant record: from the earliest plant spores to the earliest seeds, 437-477. In Talent, J. (ed.) Earth and Life - Global biodiversity, extinction intervals and biogeographic perturbations through time. Springer, Dordrecht, London & New York.
Štorch, P. 1995. Upper Silurian (upper Ludlow) graptolites of the N. inexpectatus and N. kozlowskii biozones from Kosov Quarry near Beroun (Barrandian area, Bohemia). Bulletin of the Czech Geological Survey 70, 65-89.
Štorch, P. 2023. Graptolite biostratigraphy and biodiversity dynamics in the Silurian System of the Prague Synform (Barrandian area, Czech Republic). Bulletin of Geosciences 98(1), 1-78.
Szaniawski, H. 1970. Jaw apparatuses of the Ordovician and Silurian polychaetes from the Mielnik borehole. Acta Palaeontologica Polonica 15(4), 445-472.
Talent, J.A., Mawson, R., Andrew, A.S., Hamilton, P.J. & Whitford, D.J. 1993. Middle Paleozoic extinction events: Faunal and isotopic data: Palaeo¬geography, Palaeoclimatology, Palaeoecology 104, 139-152.
Tasáryová, Z., Schnabl, P., Čížková, K., Pruner, P., Janoušek, V., Rapprich, V., Štorch, P., Manda, Š., Frýda, J. & Trubač, J. 2014. Gorstian palaeoposition and geotectonic setting of Suchomasty Volcanic Centre (Silurian, Prague Basin, Teplá-Barrandian Unit, Bohemian Massif). GFF 136, 262-265.
Tonarová, P. 2012. Silurian scolecodonts of the Barrandian area. 192 pp. Ph.D. thesis, Charles University in Prague, Czech Republic.
Tonarová, P., Eriksson, M.E. & Hints, O. 2012. A jawed polychaete fauna from the late Ludlow Kozlowskii event interval in the Prague Basin (Czech Republic). Bulletin of Geosciences 87(4), 713-732.
Tonarová, P., Hints, O. & Eriksson, M.E. 2014. Impact of the Silurian Ireviken Event on polychaete faunas: new insights from the Viki drill core, western Estonia. GFF 136(1), 270-274. ISSN 1103-5897.
Tonarová, P., Hints, O., Königshof, P., Suttner, T., Kido, E., Da Silva, A. & Pas, D. 2016. A Middle Devonian jawed polychaete fauna from the type Eifel area, western Germany, and its biogeographical and evolutionary affinities. Papers in Palaeontology 2(2), 295-310.
Tonarová, P., Vodrážková, S., Ferrová, L., de la Puente, G.S., Hints, O., Frýda, J. & Kubajko, M. 2017. Palynology, microfacies and biostratigraphy across the Daleje Event (Lower Devonian, lower to upper Emsian): new insights from the offshore facies of the Prague Basin, Czech Republic. Palaeobiodiversity and Palaeoenvironments 97, 419-438.
Tonarová, P., Vodrážková, S., Hints, O., Mannik, P., Kubajko, M. & Frýda, J. 2019. Llandovery microfossils and microfacies of the Hýskov section, Prague Basin. Fossil Imprint 75(1), 25-43.
Tonarová, P., Vodrážková, S., Hints, O., Nolvak, J., Kubajko, M. & Čáp, P. 2023. Latest Ordovician jawed polychaetes, chitinozoans and depositional environments of the Levín section, Prague Basin, Czech Republic. Geobios 81, 179-198.
Tonarová, P., Suttner, T.J., Hints, O., Liang, Y., Zemek, M., Kubajko, M., Zikmund, T., Kaiser, J. & Kido, E. 2024. Late Ordovician scolecodonts and chitinozoans from the Pin Valley in Spiti, Himachal Pradesh, northern India. Acta Palaeontologica Polonica 69(2), 199-215.
Torsvik, T.H. & Cocks, L.R.M. 2017. Earth History and Palaeogeography. 317 pp. Cambridge University Press, Cambridge & New York.
Trotter, J.A., Williams, I.S., Barnes, C.R., Männik, P. & Simpson, A. 2016. New conodont δ18O records of Silurian climate change: implications for environmental and biological events. Palaeogeography, Palaeoclimatology, Palaeoecology 443, 34-48.
Urbanek, A. 1993. Biotic crises in the history of the Upper Silurian graptoloids: a palaeobiological model. Historical Biology 7, 29-50.
Vandenbroucke, T.R.A., Hints, O., Williams, M., Wallis, S., Velleman, J., Kurihara, T., Tanaka, G., Komatsu, T., Männik, P., Siveter, D.S. & Backer, T. de 2019. Chitinozoans and scolecodonts from the Silurian and Devonian of Japan. Island Arc 28(3), 1-13.
Vecoli, M., Riboulleau, A. & Versteeghm G.J.M. 2009. Palynology, organic geochemistry and carbon isotope analysis of a latest Ordovician through Silurian clastic succession from borehole Tt1, Ghadamis Basin, southern Tunisia, North Africa: Palaeoenvironmental interpretation. Palaeogeography, Palaeoclimatology, Palaeoecology 273, 378-394.
Vodička, J. 2012. Chitinozoa nejvyššího ludlow pražské pánve a jejich význam pro korelace a vývoj faun napříč kozlowskii/lau eventem. 73 pp. Master thesis, Charles University, Prague, Czech Republic.
Vokáč, V., Hartl, F. & Pavlovič, M. 2019. Trilobiti v lomu Kosov u Berouna: Barrandien, pražská pánev, Česká republika. 52 pp. Miroslav Pavlovič, Stod.
Wenzel, B. 1997. Isotopenstratigraphische Untersuchnungen an silurischen Abfolgen und deren paläoozeanographische Interpretation. Erlanger geologischen Abhandlungen 129, 1-117.
Wenzel, B. 1999. Biostratigraphy and geochemistry of Upper Silurian brachiopods from the Timan-Pechora region (Russia). Acta Geologica Polonica 49, 145-157.
Wenzel, B. & Joachimski, M.M. 1996. Carbon and oxygen isotopic composition of Silurian brachiopods (Gotland/Sweden). Palaeoceanographic implications. Palaeogeography, Palaeoclimatology, Palaeoecology 122, 143-166.
Whalen, C.D. & Briggs, D.E.G. 2018 The Palaeozoic colonization of the water column and the rise of global nekton. Proceedings of the Royal Society B 285, 20180883.
Williams, S.H., Nowlan, G.S., Barnes, C.R. & Batten, R.S.R. 1999. The Ledge Section at Cow Head, western Newfoundland, as GSSP candidate for the lower boundary of the second stage of the Ordovician System: new data and discussion of the graptolite, conodont and chitinozoan assemblages. 33 pp. A report to the IUGS/ICS Subcommission on Ordovician Stratigraphy.
Younes, H., Calner, M. & Lehnert, O. 2017. The first continuous δ13C record across the Late Silurian Lau Event on Gotland, Sweden. GFF 139, 63-69.
Žák, J. & Sláma, J. 2018. How far did the Cadomian ?terranes’ travel from Gondwana during early Palaeozoic? A critical reappraisal based on detrital zircon geochronology. International Geology Review 60, 313-338.
Žebera, K. 1935. Les Conodontes et les Scolécodontes du Barrandien. Bulletin of international Academy of Sciences de Boheme 36, 88-96.
Zhang, F., Lenton, T.M., del Rey, Á., Romaniello, S.J., Chen, X., Planavsky, N.J., Clarkson, M.O., Dahl, T.W., Lau, K.V., Wang, W., Li, Z., Zhao, M., Isson, T., Algeo, T.J. & Anbar, A.D. 2020. Uranium isotopes in marine carbonates as a global ocean paleoredox proxy: A critical review. Geochimica et Cosmochimica Acta 287, 27-49.
Zhang, F., Frýda, J., Fakhraee, M., Lin, Y., Wei, G., Cao, M., Li, N., Zhou, J., Frýdová, B., Wei, H. & Shen, S. 2022. Marine anoxia as a trigger for the largest Phanerozoic positive carbon isotope excursion: Evidence from carbonate barium isotope record. Earth and Planetary Science Letters 584, 117421.
