Digestive structures in the Middle Ordovician trilobite Prionocheilus Rouault 1847, from the Barrandian area of Czech Republic


  • O. FATKA Charles university, faculty of Sciences, institute of Geology and Palaeontology Albertov 6, 128 43, Prague, Czech Republic.
  • P. BUDIL Czech Geological Survey Klárov 3, 118 21, Prague, Czech Republic.




Prionocheilus, Digestive tract, Ordovician, Barrandian area, Czech Republic.


Remains of a digestive system from a slightly damaged articulated specimen of the comparatively rare bathycheilid trilobite Prionocheilus vokovicensis (Šnajdr, 1956) are described for the first time. The specimen comes from the Middle Ordovician Šárka Formation of the Prague Basin and contains the midgut region of the digestive system preserved through the axial region of glabella and six anterior thoracic segments. The anterior-most part of the digestive system is unknown as the anterior glabellar lobes are not preserved in the studied specimen. In the cephalic shield, the remains of two pairs of gut diverticulae are seen in the posterior region of the glabella. Remains of five pairs of small cavities developed in the axis of the first six thoracic segments represent the remains of thoracic gut diverticulae. The discussed specimen possess the first undoubted remain of digestive structures established within the family Bathycheilidae (PŘibyl, 1953).


Apollonov, M.K. 1974. Ashgillskie trilobity Kazachstana [Ashgill trilobites from Kazakhstan]. lzdatel’stvo Nauka Kasachskoj ASSR, Alma-Ata. 136pp. [in Russian]

Aubrechtová, M., 2015. A revision of the Ordovician cephalopod Bactrites sandbergeri Barrande: Systematic position and palaeobiogeography of Bactroceras. Geobios, 48(3), 193-211. DOI: 10.1016/j.geobios.2015.03.002

Babcock, L.E., Chang, W.T., 1997. Comparative taphonomy of two nonmineralized arthropods: Naraoia (Nektaspida; Early Cambrian, Chengjiang Biota, China) and Limulus (Xiphosurida; Holocene, Atlantic Ocean). Bulletin of the National Museum of Natural Science, 10, 233-250.

Babcock, L.E., Merriam, D.F., West, R.R., 2000. Paleolimulus, an early limuline (Xiphosurida), from Pennsylvanian-Permian

Lagerstätten of Kansas, and taphonomic comparison with modern Limulus. Lethaia, 33(3), 129-141.

Barrande, J., 1846. Notice préliminaire sur le Systême Silurien et les trilobites de Bohême. C.L. Hirschfeld, libraire. Leipzig, 97pp. DOI: 10.5962/bhl.title.9141

Barrande, J., 1872. Systême Silurien du centre de la Bohême. I, Supplement au vol. 1. Trilobites, crustacés divers et poissons. Prague and Paris, 647pp.

Bergström, S.M., Chen, X., Gutiérrez-Marco, J.C., Dronov, A., 2008. The new chronostratigraphic classification of the Ordovician System and its relations to major regional series and stages and to δ13C chemostratigraphy. Lethaia, 42(1), 97-107. DOI: 10.1111/j.1502-3931.2008.00136.x

Bertling, M., 1992. Arachnostega n. ichnog. - burrowing traces in internal moulds of boring bivalves (Late Jurassic, Northern Germany). Paläontologische Zeitschrift, 66(1-2), 177-185.

Bruthansová, J., 2003. The trilobite Family Illaenidae Hawle et Corda, 1847 from the Ordovician of the Prague Basin (Czech Republic). Transactions of the Royal Society Edinburgh, Earth Sciences, 93(2), 167-190.

Budil, P., Fatka, O., 2008. Bohemian and Moravian Trilobites and their relatives. Czech Geological Survey, Prague, 47pp. ISBN: 978-80-7075-711-6.

Budil, P., Kraft, P., Kraft, J., Fatka, O., 2007. Faunal associations of the Šárka Formation (Middle Ordovician, Darriwilian, Prague Basin, Czech Republic). Acta Palaeontologica Sinica, 46(Supplement), 64-70.

Butler, A.D., Cunningham, J.A., Budd, G.E., Donoghue, P.C.J., 2015. Experimental taphonomy of Artemia reveals the role of endogenous microbes in mediating decay and fossilization. Proceedings of the Royal Society Biological Sciences, 282(1808), 20150476. DOI: 10.1098/rspb.2015.0476

Chlupáč, I., 1970. Phyllocarid crustaceans of the Bohemian Ordovician. Sborník Geologických Věd, Paleontologie, 12, 41-77.

Chlupáč, I., 2003. Phyllocarid crustaceans from the Middle Ordovician Šárka Formation at Praha-Vokovice. Bulletin of Geosciences, 78(1), 107-111.

Chvátal, M., 2003. Mineral assemblage of the Červený vrch locality. Bulletin of Geosciences, 78(2), 103-106.

Cooper, G.A., Kindle, C.H. 1936. New brachiopods and trilobites from the Upper Ordovicia of Percé, Quebec. Journal of Paleontology 10, 348-372.

Courtessole, R., Pillet, J. 1975. Contribution à l’etude des faunes trilobitiques de l’Ordovicien inférieur de la Montagne Noire. Les Eulominae et les Nileidae. Annales de la Société Géologique du Nord, 95, 251-272.

Dabbard, M.-P., Loi, A., 2012. Environmental control on concretion-forming processes: Examples from Paleozoic terrigenous sediments of the North Gondwana margin, Armorican Massif (Middle Ordovician and Middle Devonian) and SW Sardinia (Late Ordovician). Sedimentary Geology, 267-268, 93-103. DOI: 10.1016/j.sedgeo.2012.05.013

Daley, A.C., Drage, H.B., 2016. The fossil record of ecdysis, and trends in the moulting behaviour of trilobites. Arthropod

Structure & Development, 45, 71-96. DOI: 10.1016/j.asd.2015.09.004

Dean, W.T. 1964. The status of the Ordovician trilobite genera Prionocheilus and Polyeres. Geological Magazine, 101(1), 95-96.

Drost, K., Linnemann, U., Wemmer, K., Budil, P., Kraft, P., Fatka, O., Marek, J., 2003. Provenance, geotectonic setting, and early diagenetic processes of the Šárka Formation at Praha – Červený vrch (Ordovician, Barrandian, Czech Republic). Bulletin of Geosciences, 78(1), 147-156.

Fatka, O., Mergl, M., 2009. The “microcontinent” Perunica: status and story 15 years after conception. In: Bassett, M.G. (ed.).

Early Palaeozoic Peri-Gondwanan Terranes: New Insights from Tectonics and Biogeography. London, Geological Society, Special Publications, 325, 65-102. DOI: 10.1144/SP325.4

Fatka, O., Mikuláš, R., Szabad, M., Micka, V., Valent, M., 2011. Arachnostega Bertling, 1992, in Cambrian of the Barrandian

area (Czech Republic). Acta Geologica Polonica, 61(4), 367-381.

Fatka, O., Lerosey-Aubril, R., Budil, P., Rak, Š., 2013. Fossilised guts in trilobites from the Upper Ordovician Letná Formation

(Prague Basin, Czech Republic). Bulletin of Geosciences, 88(1), 95-104. DOI: 10.3140/bull.geosci.1329

Fatka, O., Budil, P., David, M., Kozák, V., Micka, V., Szabad, M., 2014. Digestive structures in Cambrian and Ordovician trilobites from the Barrandian area (Czech Republic). In: Zhan, R.B., Huang, B. (eds.). IGCP 591 Field Workshop 2014, Kunming China 12-21 August 2014. Nanjing University Press, Extended Summary, 49-51.

Fatka, O., Budil, P., David, M., 2015. Digestive structures in Ordovician trilobites Colpocoryphe and Flexicalymene from the Barrandian area of Czech Republic. Estonian Journal of Earth Sciences, 64(4), 255-266. DOI: 10.3176/earth.2015.32

Gutiérrez-Marco, J.C., García-Bellido, D.C., Rábano, I., Sá, A.A., 2017. Digestive and appendicular soft-parts, with behavioural implications, in large Ordovician trilobite from the Fezouata Lagerstätte, Morocco. Scientific Reports, 7, 39728. DOI:10.1038/srep39728

Hammann, W., 1983. Calymenacea (Trilobita) aus dem Ordovizium von Spanien; ihre Biostratigraphie, Ökologie und Systematik. Abhandlungen der Senckenbergischen naturforschenden Gesellschaft, 542, 1-177. [in German, with English abstract]

Hammann, W., Henry, J.L., 1978. Quelques éspeces de Calymenella, Eohomalonotus et Kerfornella (Trilobita, Ptychopariidae de l’Ordovicien du Massif Armoricain et de la Péninsule Ibérique). Senckenbergiana Lethaea, 59(4/6), 401-429. [in French, with English abstract]

Havlíček, V., 1981. Development of a linear sedimentary depression exemplified by the Prague Basin (Ordovician–Middle Devonian; Barrandian area – central Bohemia). Sborník Geologických Věd, Geologie, 35, 7-48.

Havlíček, V., 1982. Ordovician of Bohemia: development of the Prague Basin and its benthic communities. Sborník Geologických Věd, Geologie, 37, 103-136.

Havlíček, V., 1998. Ordovician. In: Chlupáč, I., Havlíček, V., Kříž, J., Kukal, Z., Štorch, P. (eds.). Palaeozoic of the Barrandian.

Czech Geological Survey, Prague, 149-164.

Havlíček, V., Fatka, O., 1992. Ordovician of the Prague Basin (Barrandian area, Czechoslovakia). In: Webby, B.D., Laurie, J.R. (eds.). Global perspectives on Ordovician geology. Balkema, Rotterdam, 461-472.

Havlíček, V., Vaněk, J., 1966. The Biostratigraphy of the Ordovician of Bohemia. Sborník Geologických Věd, Paleontologie, 8, 7-69.

Havlíček, V., Vaněk, J., 1990. Ordovician invertebrate communities in black-shale lithofacies (Prague Basin, Czechoslovakia). Věstník Ústředního ústavu Geologického, 65(4), 223-236.

Hawle, I., Corda, A.J.C., 1847. Prodrom einer Monographie der böhmischen Trilobiten. Abhandlungen der königlichen böhmischen Gesellschaft der Wissenschaften, 5, 119-292.

Horný, R., 2001. Ordovician Tergomya and isostrophic Gastropoda (Mollusca) of Bohemia: Types and referred specimens in the collections of the National Museum, Prague, Czech Republic. Acta Musei Nationalis Pragae, Series B – Historia Naturalis, 57(3-4), 69-102.

Horný, R., Bastl, F., 1970. Type specimens of fossils in the National Museum of Prague. Volume 1, Trilobita; [Scientific revisors: Chlupáč, I., Marek, L., Přibyl, A., Šnajdr, M.]. Prague, Museum of Natural History, 354pp.

Hupé, P., 1953. Classification des Trilobites. Annales de Paléontologie, 39, 61-168.

Ingham, J.K., 1977. A monograph of the upper Ordovician trilobites from the Cautley and Dent districts of Westmorland and Yorkshire. Palaeontographical Society Monograph, 3, 89-121.

Jaekel, O., 1901. Ueber die Organisation der Trilobiten. Zeitschrift der Deutschen Geologischen Gesseschaft, 53(1), 133-171. [in German]

Kraft, J., Kraft, P., 1992. Biostratigraphy of the Klabava and Šárka Formations (Bohemia, Lower Ordovician) – a brief overview of new investigations. Acta Universitatis Carolinae, Geologica, 33(1-2), 23-29.

Kraft, J., Kraft, P., 1999. Graptolite biozones of the Bohemian Lower and Middle Ordovician and their historical development. Journal of the Czech Geological Society, 44(1-2), 53-62.

Kraft, P., Kraft, J., 2003. Middle Ordovician graptolite fauna from Praha – Červený vrch (Prague Basin, Czech Republic). Bulletin of Geosciences, 78(1), 129-139.

Kraft, P., Kraft, J., Prokop, R.J., 2001. A possible hydroid from the Lower and Middle Ordovician of Bohemia. Alcheringa, 25(2), 143-154.

Kukal, Z., 1962. Petrografický výzkum šáreckých vrstev barrandienského ordoviku [Petrographical investigation of the Ordovician Šárka beds in the Barrandian area]. Sborník Ústředního ústavu geologického, odd. geologický, 27, 175-214. [in Czech, with English abstract]

Lajblová, K., Kraft, P., 2014. The earliest ostracods from the Ordovician of the Prague Basin, Czech Republic. Acta Geologica Polonica, 64(4), 367-392.

Lefebvre, B., 2007. Early Palaeozoic palaeobiogeography and palaeoecology of stylophoran echinoderms. Palaeogeography, Palaeoclimatology, Palaeoecology, 245, 156-199. DOI: 10.1016/j.palaeo.2006.02.021

Lemke, U., 2016. Katalog verfügbarer Namen der Trilobiten-Taxa. Unpublished, 503pp. [Available at: https://www.researchgate.net/publication/297418418_Katalog_verfugbarer_Namen_der_Trilobiten-Taxa_Stand_Marz_2016]

Lerosey-Aubril, R., Hegna, T.A., Olive, S., 2011. Inferring internal anatomy from the trilobite exoskeleton: the relationship between frontal auxiliary impressions and the digestive system. Lethaia, 44(2), 166-184. DOI: 10.1111/j.1502-


Lerosey-Aubril, R., Hegna, T.A., Kier, C., Bonino, E., Habersetzer, J., Carré, M., 2012. Controls on gut phosphatisation: The

Trilobites from the Weeks Formation Lagerstätte (Cambrian; Utah). PLOS ONE, 7(3), e32934. DOI:10.1371/journal.pone.0032934

Manda, Š., 2008. Trocholites Conrad, 1838 (Nautiloidea, Tarphycerida) in the Middle Ordovician of the Prague Basin and its palaeobiogeographical significance. Bulletin of Geosciences, 83(3), 327-334. DOI: 10.3140/bull.geosci.2008.03.327

McCoy, F. 1846. A synopsis of the Silurian fossils of Ireland. University Press, M.H. Gill, Dublin, 72pp.

Mergl, M., 2002. Linguliformean and craniiformean brachiopods of the Ordovician (Třenice to Dobrotivá formations) of the Barrandian, Bohemia. Acta Musei Nationalis Pragae, Series B, 58, 1-82.

Mergl, M., Kozák, V., 2016. The effaced Ordovician trilobite Svobodapeltis and the earliest history of illaenimorph trilobites in the Ordovician of the Prague Basin (Bohemia). Bulletin of Geosciences, 91(1), 155-168. DOI: 10.3140/bull.geosci.1516

Mergl, M., Fatka, O., Budil, P., 2007. Lower and early Middle Ordovician trilobite associations of the Prague Basin (Perunica, Czech Republic). Acta Palaeontologica Sinica, 6 (Supplement), 320-327.

Mergl, M., Fatka, O., Budil, P., 2008. Lower and Middle Ordovician trilobite associations of Perunica: from shoreface endemicicty to offshore uniformity (Prague Basin, Czech Republic). In: Rábano, I., Gozalo, R., García-Bellido, D. (eds.). Advances in trilobite research. Instituto Geológico y Minero de España, Cuadernos del Museo Geominero, 9, 275-282.

Mikuláš, R., 1991. Trace fossils from siliceous concretions in the Šárka and Dobrotivá Formations (Ordovician, central Bohemia). Časopis pro mineralogii a geologii, 36(1), 29-38.

Mikuláš, R., 1998. Ordovician of the Barrandian area: Reconstruction of the sedimentary basin, its benthic communities and ichnofacies. Journal of the Czech Geological Society, 43(3), 143-159.

Peršín, J., Budil, P., 2009. Nové poznatky ze šáreckého a dobrotivského souvrství (ordovik, stupeň darriwil) v severozápadní a severní části Prahy [New observations on Šárka and Dobrotivá Formation (Ordovician, Darriwilian Stage) in northwestern and northern surroundings of Prague]. Český kras, 35, 25-35. [in Czech, with English abstract]

Polechová, M., 2013. Bivalves from the Middle Ordovician Šárka Formation (Prague Basin, Czech Republic). Bulletin of Geosciences, 88(2), 427-461. DOI: 10.3140/bull.geosci.1426

Přibyl, A., 1953. Seznam eských trilobitových rodů. [Index of trilobite genera in Bohemia]. Knihovna Ústředního ústavu geologického, 25, 1-80. [in Czech and English]

Rábano, I., 1989. Trilobites del Ordovicico Medio del sector meridional de la Zona Centroibérica Española. III. Calymenina

y Cheirurina. Boletin Geológico y Minero, 100(5), 767-841.

Rouault, M., 1847. Extrait du Mémoire sur les Trilobites du Département d’Ille-et-Villaine. Bulletin de la Société Géologique de France, 4, 309-328.

Servais, T., Dzik, J., Fatka, O., Heuse, T., Vecoli, M., Verniers, J., 2008. Ordovician. In: McCann, T., (ed.). Geology of Central Europe. London, Geological Society, 203-248.

Siveter, D.J., 1973. Trilobites of the family Calymenidae from the Ordovician, Silurian and Devonian systems of North-West

Europe. PhD Thesis, University of Leicester, 380pp.

Siveter, D.J., 1976. The Middle Ordovician of the Oslo Region, Norway. 27. Trilobites of the family Calymenidae. Norsk Geologisk Tidsskrift, 56, 335-396.

Šnajdr, M., 1956. Trilobiti drabovských a letenských vrstev českého ordoviku. [The trilobites from the Drabov and Letná Beds of the Ordovician of Bohemia]. Sborník Ústředního ústavu geologického, 22, 477-533. [in Czech, with English abstract]

Šnajdr, M., 1990. Bohemian trilobites. Prague, Czech Geological Survey, 265pp.

Steinová, M., 2012. Probable ancestral type of actinodont hinge in the Ordovician bivalve Pseudocyrtodonta Pfab, 1934. Bulletin of Geosciences, 87(2), 333-346. DOI: 10.3140/bull.geosci.1330

Strang, K.M., Armstrong, H.A., Harper, D.A.T., 2016. Minerals in the gut: scoping a Cambrian digestive system. Royal Society

of Open Science, 3, 160420. DOI: 10.1098/rsos.160420

Valíček, J., Vaněk, J., 2001. New index of the genera, subgenera, and species of Barrandian trilobites. Part A-B (Cambrian and Ordovician). Palaeontologia Bohemiae, 7, 1-49.

Vaněk, J., 1965. New species of the suborder Calymenina Swinneton, 1915 (Trilobita) from the Barrandian area. Sborník geologických věd, Paleontologie, 6, 21-37.

Vaněk, J., 1995. New deeper water trilobites in the Ordovician of the Prague Basin, Czech Republic. Palaeontologia Bohemiae, 5, 1-12.

Vaněk, J., Valíček, J., 2002. New index of the genera, subgenera, and species of Barrandian trilobites. Part C-D (Silurian and

Devonian). Palaeontologia Bohemiae, 8, 1-74.

Vokáč, V., Hartl, F., David, M., Pavlovi, Doubrava, M., Kozák, V., Grigar, L., 2015. Význané nálezy trilobitů z ordoviku (daping–sandby) pražské pánve (Barrandien, Česká Republika) (Notable findings of trilobites from the Ordovician (Dapingian–Sandbian) of the Prague Basin (Barrandian area, Czech Republic). Erica, 22, 141-157. [in Czech, with English abstract].

Whittington, H.B., 1965. A monograph of the Ordovician trilobites of the Bala area, Merioneth. Palaeontographical Society Monograph, 2, 33-62.

Whittington, H.B., Kelly, S.R.A., 1997. Morphological terms applied to trilobites. In: Moore, R.C., Kaesler, R.L. (eds.). Treatise on Invertebrate Paleontology, Part O Arthropoda 1 Trilobita, Revised 1. Kansas, Boulder, Colorado and Lawrence, O313-O329.

Zhou, Z.Y., Yin, G.Z., Tripp, R.P., 1984. Trilobites from the Ordovician Shihtzupu Formation, Zunyi, Guizhou Province, China. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 75(1), 13-36. DOI: 10.1017/S0263593300009755

Zhu, X.J., Lerosey-Aubril, R., Esteve, J., 2014. Gut content fossilization and evidence for detritus feeding habits in an enrolled trilobite from the Cambrian of China. Lethaia, 47(1), 66-76. DOI: 10.1111/let.12041