Response of organophosphatic brachiopods to the mid-Ludfordian (late Silurian) carbon isotope excursion and associated extinction events in the Prague Basin (Czech Republic)


Authors: Mergl M, Frýda J, Kubajko M

Published in: Bulletin of Geosciences, volume 93, issue 3; pages: 369 - 400; Received 15 April 2018; Accepted in revised form 3 July 2018; Online 6 August 2018

Keywords: mid-Ludfordian carbon isotope excursion, Lau and Kozlowskii bioevents, Silurian, organophosphatic brachiopods, Barrandian, new taxa,

full text (PDF, 12.77 MB)

Export to RIS



The present paper represents the first results of an on-going multidisciplinary study focused on the response of marine faunas to the mid-Ludfordian (late Silurian) carbon isotope excursion and associated extinction events (Ludfordian, late Silurian). The paper describes the stratigraphical distribution of organophosphatic brachiopods from the uppermost part of the Neocucullograptus inexpectatus to lower part of Monograptus parultimus graptolite biozones (i.e. early Ludfordian to earliest Přídolí). Numerical analysis of the stratigraphical distribution, based on more than 1300 determinable valves of 15 organophosphatic brachiopod species, revealed three groups being significantly supported. They are here formalized as new organophosphatic brachiopod communities: the oldest Opsiconidion ephemerus Community, the Kosoidea fissurella Community, and the youngest Opsiconidion parephemerus-Kosagittella clara Community. The Opsiconidion ephemerus Community and the Opsiconidion parephemerus-Kosagittella clara Community have distinct similarities in their taxonomic composition, moderate species diversity and low dominance indices. The communities are separated by the monospecific Kosoidea fissurella Community present just after the Lau and Kozlowskii bioevents during the period of high δ13C values which is interpreted as an opportunistic community. Re-occurrence of some brachiopod taxa after the bioevents and the mid-Ludfordian carbon isotope excursion raises the question, at least for organophosphatic brachiopods, as to the extent to which the Lau and Kozlowskii bioevents represent true global extinction events and whether their significance has not been overestimated. Two new species are erected: Opsiconidion bouceki and Opsiconidion parephemerus.


Andrew, A.S., Hamilton, P.J., Mawson, R., Talent, J.A. & Whitford, D.J. 1994. Isotopic correlation tools in the mid-Paleozoic and their relation to extinction events. APEA Journal 34, 268-77.

Baliński, A. & Holmer, L.E. 1999. The Late Devonian trematid lingulate brachiopod Schizobolus from Poland. Acta Palaeontologica Polonica 44, 335-346.

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, 13-33.

Bassett, M.G. 1984. Life strategies of Silurian brachiopods. Special Papers in Palaeontology 32, 237-263.

Bassett, M.G., Popov, L.E., Aldridge, R.J., Gabbott, S.E. & Theron, J.N. 2009. Brachiopoda from the Soom Shale Lagerstätte (upper Ordovician, South Africa). Journal of Paleontology 83, 614-623. View article

Biernat, G. & Harper, D.A.T. 1999. A lingulate brachiopod Acrotretella: New data from Ordovician of Poland. Acta Palaeontologica Polonica 44, 83-92.

Botting, J.P. 2002. The role of pyroclastic volcanism in Ordovician diversification, 99-113. In Crame, J.A. & Owen, A.W. (eds). Palaeobiogeography and biodiversity change: the Ordovician and Mesozoic- Cenozoic radiations. Geological Society London, Special Publication 194.

Bouček, B. 1934. Bemerkungen zur Stratigraphie des böhmischen Gotlandien und seinen Faziesverhaltnissen. Centralblatt für Mineralogie, Geologie und Palaeontologie, Abt. B 11, 477-494.

Bouček, B. & Přibyl, A. 1955. O silurských ostrakodech a stratigrafii vrstev budňanských z nejbližšího okolí Kosova a Koledníku u Berouna. Sborník Ústředního ústavu geologického, Oddíl paleontologický 21, 577-662.

Boucot, A.J. 1975. Evolution and extinction rate controls. 427 pp. Elsevier, Balkema.

Boucot, A.J., Rowell, A.J., Racheboeuf, P., Pereira, E., Gonçalves de Melo, J.H. & Peixoto de Siqueira, L. 2001. Position of the Malvinokaffric Realm’s northern boundary (Early Devonian) based on newly discovered brachiopods from the Parecis Basin (Brazil). Journal of the Czech Geological Society 46, 109-120.

Brett, C.E., Ferretti, A., Histon, K. & Schonlaub, H.P. 2009. Silurian sequence stratigraphy of the Carnic Alps, Austria. Palaeogeography, Palaeoclimatology, Palaeoecology 279, 1-28. View article

Brock, G.A., Engelbretsen, M.J. & Dean-Jones, G. 1995. Acrotretoid brachiopods from the Devonian of Victoria and New South Wales. Memoirs of the Association of Australasian Palaeontologists 18, 105-120.

Calner, M. 2008. Silurian global events - at the tipping point of climate change, 21-58. In Ashraf, M.T. (ed.) Mass extinctions. Springer-Verlag. Berlin & Heidelberg.View article

Cherns, L. 1979. The environmental significance of Lingula in the Ludlow Series of the Welsh Borderland and Wales. Lethaia 12, 35-46. View article

Clarke, J.M. 1913. Fósseis Devonianos do Paraná. Monographias do Servico Geológico e Mineralógico do Brasil 1, 1-353.

Cocks, L.R.M. 1979. New acrotretacean brachiopods from the Palaeozoic of Britain and Austria. Palaeontology 22, 93-100.

Cocks, L.R.M. & Popov, L. 2009. Generic homes for British linguloid brachiopods. Palaeontology 52, 349-367. View article

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. View article

Cooper, G.A. 1956. Chazyan and related brachiopods. Smithsonian Miscellaneous Collection 127, 1-1245.

Duméril, A.M.C. 1806. Zoologie analytique ou méthode naturelle de classi?cation des animaux. xxiv + 344 pp. Allais, Paris.

Ebbestad, J.O.R., Frýda, J., Wagner, P., Horný, R., Isakar, M., Stewart, S., Percival, I., Bertero, V., Rohr, D.M., Peel, J.S., Blodgett, R.B. & Högström, A.E.S. 2013. Biogeography of Ordovician and Silurian gastropods, monoplacophorans and mimospirids, 191-212. In Harper, D.A.T. & Servais, T. (eds) Early Palaeozoic Biogeography and Palaeogeography. Geological Society London, Memoir 38(1). View article

Emig, C.C. 1997. Ecology of inarticulated brachiopods, 473-495. In Williams, A., Brunton, C.H.C., Carlson, S. J. et al. (eds) Treatise on Invertebrate Paleontology, part H, Brachiopoda, Revised. Geological Society of America Inc. and The University of Kansas, Boulder, Lawrence.

Eriksson, M.J., Nilsson, E.K. & Jeppsson, L. 2009. Vertebrate extinctions and reorganizations during the Late Silurian Lau Event. Geology 37, 739-742. View article

Farkaš, J., Frýda, J. & Holmden, C. 2016. Calcium isotope constraints on the marine carbon cycle and CaCO3 deposition during the late Silurian positive δ13C excursion. Earth and Planetary Science Letters 451, 31-40. View article

Farkaš, J., Frýda, J. & Holmden, C. 2017. Corrigendum: Calcium isotope constraints on the marine carbon cycle and CaCO3 deposition during the late Silurian positive δ13C excursion. Earth and Planetary Science Letters 469, 170-171. View article

Frýda, J. & Frýdová, B. 2017. First paired carbonate-carbon and organic carbon isotope records across the Ludfordian (late Silurian) suggest elevated pCO2 before the largest Phanerozoic carbon isotope anomaly. GSA Annual Meeting in Seattle, Washington, USA, Abstract ID# 299437.

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. View article

Gocke, M., Lehnert, O. & Frýda, J. 2013. Facies development and palynomorphs during the Lau Event (Late Silurian) in a non-tropical carbonate environment: shallow and deep water examples from the Barrandian Area (Czech Republic). Facies 59, 611-630. View article

Gray, J.E. 1840. Synopsis of the contents of the British Museum, 42th edition. 370 pp. British Museum, London.

Harper, D.A.T. 1984. Brachiopods from the Upper Ardmillan succession (Ordovician) of the Girvan district, Scotland. Part 1. Palaeontographical Society Monograph 136, 1-78.

Havlíček, V. & Mergl, M. 1988. Two new discinid genera (Brachiopoda) from the Silurian and Devonian of the Prague Basin, Czechoslovakia. Věstník Českého geologického ústavu 63, 169-172.

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. 1996. Brachiopods and trilobites in the Chýnice Limestone (Emsian) at Bubovice (Čeřinka hillside; Prague Basin). Palaeontologica Bohemiae 2, 1-16.

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, Geologie 46, 23-56.

Hiller, N. 1993. A modern analogue for the Lower Ordovician Obolus conglomerate of Estonia. Geological Magazine 130, 365-367. View article

Histon, K. 2012 The Silurian nautiloid-bearing strata of the Cellon Section (Carnic Alps, Austria): Color variation related to events. Palaeogeography, Palaeoclimatology, Palaeoecology 367-368, 231-255. View article

Histon, K. & Schönlaub, H. P. 1999. The Palaeozoic of the Southern Alps. Berichte der Geologisches Bundesanstalt 47, 6-30.

Holmer, L.E. 1986. Inarticulate brachiopods around the Middle-Upper Ordovician boundary in Västergötland. Geologiska Föreningens i Stockholm Förhandlingar 108, 97-126. View article

Holmer, L.E. 1989. Middle Ordovician phosphatic inarticulate brachiopods from Västergötland and Dalarna, Sweden. Fossils and Strata 26, 1-172.

Holmer, L.E. & Popov, L.E. 2000. Lingulata, 30-146. In Williams, A., Brunton, C.H.C. & Carlson, S.J. et al. (eds) Treatise on Invertebrate Paleontology, part H, Brachiopoda, Revised, Volume 2. Geological Society of America Inc. and The University of Kansas, Boulder, Lawrence.

Horný, R. 1955. Studie o vrstvách budňanských v západní části Barrandienu. Sborník Ústředního ústavu geologického, Oddíl geologický 21, 315-447.

Ireland, H.A. 1961. New phosphatic brachiopods from the Silurian of Oklahoma. Journal of Paleontology 35, 1137-1142.

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.

Jeppsson, L., Talent, J.A., Mawson, R., Andrew, A., Corradini, C., Simpson, A.J., Wigforss-Lange, J & Schönlaub, A.P. 2012. Late Ludfordian correlation and the Lau Event. In Talent, J.A. (ed.) Earth and Life, International Year of Planet Earth. Springer Science 2012.View article

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. View article

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, 211-223. View article

Kaljo, D., Boucot, A.J., Corfield, R.M., Le Herisse, A., 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.View article

Kaljo, D., Grytsenko, V., Kallaste, T., Kiipli, T. & Martma, T. 2014. Upper Silurian stratigraphy of Podolia revisited: carbon isotopes, bentonites and biostratigraphy. Geologiska Föreningens i Stockholm Förhandlingar 136, 136-141. View article

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(4), 205-220. View article

King, W. 1846. Remarks on certain genera belonging to the class Palliobranchiata. Annals and Magazine of Natural History (Series 1) 18, 26-42.View article

Klyza, J.S. 1997. The Tamworth Belt in the Tamworth-Attunga area: stratigraphy, structure, biochronologic and palaeoenvironmental analysis. 238 pp. Master thesis. Macquarie University, Sydney, Australia.

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. View article

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. View article

Kříž, J. 1987. Silur, 26-34. In Chlupáč, I. (ed.) Vysvětlivky k základní geologické mapě ČSSR. 1:25 000, Králův Dvůr. Ústřední ústav geologický, Praha.

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, 179-203.

Kříž, J. 1992. Silurian field excursions: Prague Basin (Barrandian), Bohemia. National Museum of Wales, Geological Series 13, 1-111.

Kříž, J., 1998a. Silurian, 79-101. In Chlupáč, I., Havlíček, V., Kříž, J., Kukal, Z. & Štorch, P. (eds) Paleozoic of the Barrandian (Cambrian to Devonian). Český geologický ústav, Praha.

Kříž, J. 1998b. Recurrent Silurian-Lowest Devonian cephalopod limestones of Gondwanan Europe and Perunica. New York State Museum Bulletin 491, 183-198.

Kříž, J. 1999. Bivalvia dominated communities of Bohemian type from the Silurian and Lower Devonian carbonate facies, 225-248. 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.

Kříž, J. 2008. A new bivalve community from lower Ludlow of the Prague Basin. Bulletin of Geosciences 83(3), 237-280. View article

Kuhn, O. 1949. Lehrbuch der Paläozoologie. 326 pp. Schweizerbart, Stuttgart.

Kutorga, S.S. 1846. Ueber das silurische und devonische Schichten-System von Gatschina. Russisch-Kaiserliche Mineralogische Gesellschaft zu St. Petersbourg, Verhandlungen 1845-1846, 85-139.

Kutorga, S.S. 1848. Ueber die Brachiopoden-familie der Siphonotretaceae. Russisch-Kaiserliche Mineralogische Gesellschaft zu St. Petersbourg, Verhandlungen 1847, 250-286.

Lehnert, O., Frýda, J., Buggisch,W. & Manda, Š. 2003. A first report of the Ludlovian Lau event from the Prague Basin (Barrandian, Czech Republic). Serie Correlación Geológica 18, 139-144.

Lehnert, O., Eriksson, M.J., Calner, M., Joachimski, M. & Buggisch, W. 2007b. Concurrent sedimentary and isotopic incications 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. 2007a. δ13C record across the Ludlow Lau Event: new data from mid palaeo-latitudes of northern peri-Gondwana (Prague Basin, Czech Republic). Palaeogeography, Palaeoclimatology, Palaeoecology 245, 227-244. View article

Loydell, D.K. 2007. Early Silurian positive δ13C excursions and their relationship to glaciations, sea-level changes and extinction events. Geological Journal 42(5), 531-546. View article

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. View article

Ludvigsen, R. 1974. A new Devonian acrotretid (Brachiopoda, Inarticulata) with unique protegular ultrastructure. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 3, 133-148.

Manda, Š. 2008. Palaeoecology and palaeogeographic relations of the Silurian phragmoceratids (Nautiloidea, Cephalopoda) of the Prague Basin (Bohemia). Bulletin of Geosciences 83, 39-62. View article

Manda, Š. & Kříž, J. 2006. Environmental and biotic changes of the subtropical isolated carbonate platforms during Kozlowskii and Lau events (Prague Basin, Silurian, Ludlow). Geologiska Föreningens i Stockholm Förhandlingar 128, 161-168.

Manda, Š., Štorch, P., Slavík, L., Frýda, J., Kříž, J. & Tasáryová, Z. 2012. Graptolite, conodont and sedimentary record through the late Ludlow Kozlowskii Event (Silurian) in shale dominated succession of Bohemia. Geological Magazine 149, 507-531. View article

Mendez-Alzola, R. 1938. Fósiles Devónicos del Uruguay. Instituto de Geología y Perforaciones, Boletín 24, 1-115.

Mendez-Alzola, R. & Sprechmann, P. 1971. Algunas orbiculoideas devonicas de la Precordillera de San Juan, Rep. Argentina. Revista de la Asociació Geológica Argentina 24, 517-525.

Menke, C.T. 1828. Synopsis methodica molluscorum generum omnium et specierum earum quae in Museo Menkeano adservantur. 91pp. G. Uslar, Pyrmonti.

Mergl, M. 1982. Caenotreta (Inarticulata, Brachiopoda) in the Upper Silurian of Bohemia. Věstník Ústředního ústavu geologického 57, 115-116.

Mergl, M. 1999. Genus Paterula (Brachiopoda) in Ordovician Silurian sequence of Central Bohemia. Věstník Českého geologického ústavu 74, 347-361.

Mergl, M. 2001a. Lingulate brachiopods of the Silurian and Devonian of the Barrandian. Acta Musei nationalis Pragae, Series B - Historia naturalis 57, 1-49.

Mergl, M. 2001b. Extinction of some lingulate brachiopod families: new stratigraphical data from Silurian and Devonian from Bohemia, 345-351. In Brunton, C.H.C., Cocks, L.R.M. & Long, S. (eds) Brachiopods past and Present. Systematic Association Special Volume 63.

Mergl, M. 2003. Orbaspina chlupaci sp. nov., a new siphonotretid brachiopod from the Silurian of the Barrandian area, Bohemia. Bulletin of Geosciences 78, 419-421.

Mergl, M. 2009. Lingulate brachiopods from the Acanthopyge Limestone (Eifelian) of the Barrandian, Czech, Republic. Bulletin of Geoscinces 83, 281-298.

Mergl, M. 2010a. A review of Silurian discinoid brachiopods from Gotland, Sweden. Bulletin of Geosciences 85, 367-384. View article

Mergl, M. 2010b. Discinoid brachiopod life assemblages: Fossil and extant. Bulletin of Geosciences 85, 27-38. View article

Mergl, M. 2012. Lingulate and craniate brachiopods from the top of the Králův Dvůr Formation (latest Katian) and their contribution to palaeogeography. Acta Musei Nationalis Pragae, Series B - Historia Naturalis 68, 35-46.

Mergl, M. & Ferrová, L. 2009. Lingulate brachiopods from the Chýnice Limestone (upper Emsian, Barrandian; Czech Republic). Bulletin of Geosciences 84, 525-546. View article

Mergl, M. & Jiménez-Sánchez, A. 2015. Lingulate brachiopods from the Suchomasty Limestone (upper Emsian) of the Barrandian, Czech Republic. Bulletin of Geosciences 90, 173-193. View article

Mergl, M. & Massa, D. 2005. A new giant discinoid brachiopod from the Lower Devonian of Algeria. Acta Palaeontologia Polonica 50, 397-402.

Mergl, M. & Vodrážková, S. 2012. Emsian-Eifelian lingulate brachiopods from the Daleje-Třebotov Formation (Třebotov and Suchomasty limestones) and the Choteč Formation (Choteč and Acanthopyge limestones from the Prague Basin; the Czech Republic. Bulletin of Geoscinces 87, 315-332. View article

Mergl, M., Kraft, J. & Kraft, P. 2007. Life habit and spatial distribution of siphonotretid brachiopods in the Lower Ordovician of the Prague Basin, Czech Republic. Earth and Environmental Transactions of the Royal Society of Edinburgh 98, 253-261. View article

Montecino, V. & Lange, C.B. 2009. The Humboldt Current System: Ecosystem components and precesses, fisheries, and sediment studies. Progress in Oceanography 83, 65-79. View article

Morris, J. 1849. Note on the genus Siphonotreta, with a description of a new species (S. anglica). Annals and Magazine of Natural History, Series 2 & 4, 315-321. View article

Morris, J. & Sharpe, D. 1846. Description of eight species of brachiopod shells from the Paleozoic rocks of the Falkland Islands. Quarterly Journal of the Geological Society of London 2, 274-278. View article

Morzadec, P.M., Mergl, M., Villarroel, C., Janvier, P. & Racheboeuf, P.R. 2015. Trilobites and inarticulate brachiopods from the Devonian Floresta Formation of Colombia: a review. Bulletin of Geosciences 90, 331-358. View article

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, 99-124. View article

Orbigny, A. d’1847. Considérations zoologiques et géologiques sur les Brachiopodes ou Palliobranches. Comptes Rendus Hebdomodaires des Séances de l’Académie des Sciences 25, 193-195, 266-269.

Popov, L.E. 1981. Pervaia nakhodka mikroskopicheskikh bezzamkovykh brakhiopod semeistva Acrotretidae v Silure Estonii [The ?rst record of microscopic inarticulate brachiopods of the family Acrotretidae from the Silurian of Estonia]. Eesti NSV Teaduste Akadeemia Toimetised (Keemia, Geoloogia) 30, 34-41.

Popov, L.E. 2000. Late Ordovician linguliformean microbrachiopods from north-central Kazakhstan. Alcheringa 24, 257-275. View article

Popov, L.E., Nolvak, J. & Holmer, L.E. 1994. Late Ordovician lingulate brachiopods from Estonia. Palaeontology 37, 627-650.

Popov, L.E., Bassett, M.G., Holmer, L.E. & Ghobadi Pour, M. 2009. Early ontogeny and soft tissue preservation in siphonotretide brachiopods: new data from the Cambrian-Ordovician of Iran. Gondwana Research 16, 151-161. View article

Reed, F.R.C. 1925. Revision of the fauna of the Bokkeveld Beds. Annals of the South African Museum 22, 27-225.

Rowell, A.J. 1965. Inarticulata, 260-296. In Moore, R.C. (ed.) Treatise on Invertebrate Paleontology, Part H. Brachiopoda. The geological Society of America and The University of Kansas Press Boulder, Lawrence.

Satterfield, I.R. & Thompson, T.L. 1969. Phosphatic inarticulate brachiopods from the Bainbridge Formation (Silurian) of Missouri and Illinois. Journal of Paleontology 43, 1042-1048.

Schuchert, C. 1893. Classification of the Brachiopoda. American Geologist 11, 141-167.

Sharpe, D. 1856. Description of Paleozoic Mollusca from South Africa. Transactions of the Geological Society of London, Series 2, 7.

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. View article

Stampfli, G.M., Raumer, J.F & Borel, G.D. 2002. Paleozoic evolution of pre-Variscan terranes: From Gondwana to the Variscan collision. Geological Society of America Special Paper 364, 263-280.

Stricanne, L., Munnecke, A. & Pross, J. 2006. Assessing mechanisms of environmental change: Palynological signals across the late Ludlow (Silurian) positive isotope excursion (δ13C, δ18O) on Gotland, Sweden. Palaeogeography, Palaeoclimatology, Palaeoecology 230, 1-31. View article

Sutton, M.D., Bassett, M.G. & Cherns, L. 2000. Lingulate brachiopods from the Lower Ordovician of the Anglo-Welsh Basin. Part 2. Monograph of the Palaeontographical Society London 613, 61-114.

Talent, J.A., Mawson, R., Andrew, A.S., Hamilton, P.J. & Whitford, D.J. 1993. Middle Palaeozoic extinction events: Faunal and isotopic data. Palaeogeography, Palaeoclimatology, Palaeoecology 104, 139-152. View article

Tapanila, L. & Holmer, L.E. 2006. Endosymbiosis in Ordovician-Silurian corals and stromatoporoids: A new lingulid and its trace from Eastern Canada. Journal of Paleontology 88, 750-759. View article

Tasáryová, Z., Janoušek, V. & Frýda, J. 2018. Failed Silurian continental rifting at the NW margin of Gondwana - evidence from basaltic volcanism of the Prague Basin (Teplá-Barrandian Unit, Bohemian Massif). International Journal of Earth Sciences 107, 1231-1266. View article

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, 713-732. View article

Ulrich, E.O. 1886. Descriptions of new Silurian and Devonian fossils. American Paleontology, Contributions 1, 3-35.View article

Urbanek, A. 1993. Biotic crises in the history of Upper Silurian graptoloids: A palaeobiological model. Historical Biology 7, 29-50. View article

Valentine, J.L. 2006a. Early Devonian brachiopods from the Cobar Supergroup, western New South Wales, 192-259. In Valentine, J.L. (ed.) Taxonomic assessment, biostratigraphy and faunal turnover of Silurian-early Devonian linguliformean brachiopods from New South Wales, Australia. Ph.D. thesis. Macquarie University, Sydney, Australia.

Valentine, J.L. 2006b. A thorny problem revisited: a cladistic analysis of the Siphonotretida (Linguliformea, Brachiopoda), 261-342. In Valentine, J.L. (ed.) Taxonomic assessment, biostratigraphy and faunal turnover of Silurian-early Devonian linguliformean brachiopods from New South Wales, Australia. Ph.D. thesis. Macquarie University, Sydney, Australia.

Valentine, J. & Brock, G.A. 2003. A new siphonotretid brachiopod from the Silurian of Central-Western New South Wales, Australia. Records of the Australian Museum 55, 231-244. View article

Valentine, J.L., Brock, G.A. & Molloy, P.D. 2003. Linguliformean brachiopod faunal turnover across the Ireviken Event (Silurian) at Boree Creek, central-western Now South Wales, Australia, 301-327. In Königshof, P. & Schindler, E. (eds) Mid-Palaeozoic bio- and geodynamics: the north Gondwana-Laurasian interaction. Courier Forschungsinstitut Senckenberg 242.

Valentine, J.L., Cole, D. & Simpson, A.J. 2006. Silurian linguliformean brachiopods and conodonts from the Cobra Formation, Southeastern New South Wales, Australia. Proceedings of the Linnean Society of New South Wales 127, 199-234.

Vanuxem, L. 1842. Geology of New York. Part 3, comprising the survey of the third geological district, Natural History of New York (4, 3). 306 pp. D. Appleton & Co., New York.

Waagen, W. 1885. Salt Range fossils, I. Productus-Limestone fossils, Brachiopoda. Memoirs of the Geological Survey of India, Palaeontologica Indica (series 13) 4, 729-770.

Willard, B. 1928. The brachiopods of the Ottosee and Holston formations of Tennessee and Virginia. Bulletin of the Harvard Museum of Comparative Zoology 68, 255-292.

Williams, S.H. & Lockley, M.G. 1983. Ordovician inarticulate brachiopods from graptolitic shales at Dob’s Linn, Scotland; their morphology and signi?cance. Journal of Paleontology 57, 391-400.

Williams, A., Holmer, L.E. & Cusack, M. 2004. Shemico-structure of the organophosphatic shells of siphonotretide brachiopods. Palaeontology 47(5), 1313-1337. View article

Williams, A., Carlson, S.J., Brunton, C.H.C., Holmer, L. & Popov, L.E. 1996. A supra-ordinal classification of the Brachiopoda. Philosophical Transactions of the Royal Society, Biological Sciences 355, 1171-1193. View article

Wright, A.D. & McClean, A.E. 1991. Microbrachiopods and the end-Ordovician event. Historical Biology 5, 123-129. View article

Wright, A.D. & Nolvak, J. 1997. Functional significance of the spines of the Ordovician lingulate brachiopod Acanthambonia. Palaeontology 40, 113-119.

Younes, H., Calner, M. & Lehnert, O. 2016. The first continuous δ13C record across the Late Silurian Lau Event on Gotland, Sweden. Geologiska Föreningens i Stockholm Förhandlingar 1, 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, 319-338. View article

Zettler, M.L., Bochert, R. & Pollehne, F. 2009. Macrozoobenthos diversity on an oxygen minimum zone off northern Namibia. Marine Biology 156, 1949-1961. View article

Zezina, O.N. 2008. Biogeography of the Recent Brachiopods. Paleontological Journal 42, 830-858. View article