Late Bashkirian (Early Pennsylvanian) horseshoe crabs (Chelicerata: Xiphosura) from the Mospyne Formation, Donets Basin, Ukraine
Journal cover Geologos, volume 32, year 2026, title Open Volume
PDF

Keywords

Xiphosura
Belinurus
Euproops
late Bashkirian
palaeoecology

How to Cite

Dernov, V. (2026). Late Bashkirian (Early Pennsylvanian) horseshoe crabs (Chelicerata: Xiphosura) from the Mospyne Formation, Donets Basin, Ukraine. Geologos, 32. Retrieved from https://pressto.amu.edu.pl/index.php/logos/article/view/56419

Abstract

The Bashkirian xiphosurid assemblage of the Donets Basin comprises only three genera: Euproops, Prestwichianella, and Belinurus. With the exception of Euproops danae, all recorded species appear to be endemic to the basin. A small xiphosurid assemblage consisting of Belinurus sp., ?Euproops sp., and indeterminate xiphosurid remains is described from non-marine deposits of the Mospyne Formation (upper Bashkirian, Lower Pennsylvanian) in the central Donets Basin, eastern Ukraine. Together with Belinurus iswarinensis (Chernyshev, 1927), previously reported from the Bashkirian Mandrykyne and Mospyne formations, these specimens represent the oldest known Carboniferous xiphosurids from the Donets Basin.

PDF

References

Aisenverg D.Ye., Belenko N.G., Dedov V.S., Levenshtein M.L., Makarov I.A., Nesterenko L.P., Poletaev V.I., Popov V.S., Sokolova G.U., Fissunenko O.P., & Shchegolev A.K., 1975. Stratigraficheskaya ekskursiya [Stratigraphic excursion]. [In:] Aisenverg D.Ye., Lagutina V.V., Levenshtein M.L. & Popov V.S. (Eds), Putevoditel’ ekskursii po Donetskomu basseynu [Field excursion guidebook for the Donets Basin]. Nauka, Moscow, 201–245.

Aisenverg D.Ye., Brazhnikova N.E., Novik K.O., Rotai A.P. & Shulga P.L., 1963. Stratigrafiya kamennougol'nykh otlozheniy Donetskogo basseyna [Carboniferous stratigraphy of the Donets Basin]. Publishing House of the Academy of Sciences of the Ukrainian SSR, Kyiv, 182 pp.

Ambrose T. & Romano M., 1972. New Upper Carboniferous Chelicerata (Arthropoda) from Somerset, England. Palaeontology 15, 569–578.

Anderson L.I., 1994. Xiphosurans from the Westphalian D of the Radstock Basin, Somerset Coalfield, the South Wales Coalfield and Mazon Creek, Illinois. Proceedings of the Geologists’ Association 105, 265–275.

Anderson L.I., Dunlop J.A., Horrocks C.A., Winkelmann H.M. & Eagar R.M.C., 1997. Exceptionally preserved fossils from Bickershaw, Lancashire, UK (Upper Carboniferous, Westphalian A (Langsettian)). Geological Journal 32, 197–210.

Anderson L.I., Dunlop J.A., Eagar R.M.C., Horrocks C.A. & Wilson H.M., 1999. Soft-bodied fossils from the roof shales of the Wigan Four Foot coal seam, Westhoughton, Lancashire, UK. Geological Magazine 135, 321–329.

Babcock L.E. & Merriam D.F., 2000. Horseshoe crabs (Arthropoda; Xiphosurida) from the Pennsylvanian of Kansas and elsewhere. Transactions of the Kansas Academy of Science 103, 76–94.

Baily W.H., 1863. Remarks on some Coal Measures Crustacea belonging to the genus Belinurus König, with description of two new species from Queen’s County, Ireland. Annals and Magazine of Natural History 11, 107–114.

Betekhtina O.A., 1974. Biostratigrafiya i korrelyatsiya uglenosnykh otlozheniy pozdnego paleozoya po nemorskim dvustvorkam [Non-marine bivalves, biostratigraphy and correlation of the Late Paleozoic Coal Measures]. Nauka, Novosibirsk, 178 pp.

Bicknell R.D.C. & Pates S., 2020. Pictorial atlas of fossil and extant horseshoe crabs, with focus on Xiphosurida. Frontiers in Earth Science 8, 1–60.

Bicknell R.D.C, Pates S. & Botton M.L., 2019. Euproops danae (Belinuridae) cluster confirms deep origin of gregarious behaviour in xiphosurids. Arthropoda Selecta 28, 549–555.

Bicknell R.D.C., Tashman J.N., Edgecombe G.D. & Paterson J.R., 2021. Carboniferous horseshoe crab musculature suggests anatomical conservatism within Xiphosurida. Papers in Palaeontology 8, 1–9.

Bölsche W., 1885. Über Prestwichia rotundata H. Woodw. sp. aus der Steinkohlenformation des Piesberges bei Osnabrück. Jahresbericht des Naturwissenschaftlichen Vereins Osnabrück 6, 268–271.

Boyarina N.I., 2016. Makrofloristicheskiye zony srednego i verkhnego karbona (pensil'vanskoy podsistemy) Donetskogo basseyna [Macrofloristic zones of the middle and Upper Carboniferous (Pennsylvanian Subsystem) of the Donets Basin]. Geologičnij žurnal 354, 21–35.

Brauckmann C., 1982. Der Schwertschwanz Euproops (Xiphosurida, Limulina, Euproopacea) aus dem Ober-Karbon des Piesbergs bei Osnabrück. Osnabrücker Naturwissenschaftliche Mitteilungen 9, 17–26.

Bronn H.G., 1839. Fossile Limulus-Arten. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefaktenkunde 1839, 489–490.

Buckland W., 1837. The Bridgewater treatises on the power, wisdom and goodness of God as manifested in the creation. Treatise IV. Geology and mineralogy with reference to natural theology (second edition). William Pickering, London, 316 pp.

Chernyshev B.I., 1927. Zametka o predstavitelyakh Xiphosura iz Donetskogo basseyna [A note on representatives of Xiphosura from the Donets Basin]. Proceedings of the Geological Committee 46, 645–655.

Chernyshev B.I. 1928. Yeshche o Phyllopoda i Xiphosura Donetskogo basseyna [More on Phyllopoda and Xiphosura of the Donets Basin]. Proceedings of the Geological Committee 47, 519–533.

Chernyshev B.I., 1931. Carbonicola, Anthracomya i Najadites Donetskogo basseina [Carbonicola, Anthracomya, and Najadites of the Donets Basin]. Geological Publishing House of the Main Geological Prospecting Directorate, Moscow and Leningrad, 126 pp.

Chernyshev B.I., 1939. Rakoobraznye [Crustaceans]. [In:] Gorsky, I.I. (Ed.), Atlas rukovodyaschikh form iskopayemykh faun SSSR. Tom 5. Sredniy i verkhniy karbon [Atlas of index fossils of the USSR. Volume 5. Middle and Upper Carboniferous]. Gosgeoltoptekhizdat, Moscow, 141–144.

Cockerell T.D.A., 1905. Two Carboniferous genera of xiphosurans. American Geologist 36, 330.

Dernov V., 2019a. Taphonomy and paleoecology of fauna and flora from deltaic sandstones of the Mospinka Formation (middle Carboniferous) of the Donets Basin. Geo&Bio 18, 37–63.

Dernov V.S., 2019b. K izucheniyu nemorskoi fauny mospinskoi svity (sredniy karbon, Donbass) [On the study of the non-marine fauna of the Mospino Formation (Middle Carboniferous, Donets Basin)]. Tectonics and Stratigraphy 46, 105–115.

Dernov V., 2021. The earliest insect endophytic oviposition (Early Pennsylvanian, eastern Ukraine). Visnyk of the Taras Shevchenko National University of Kyiv. Geology 95, 16–24.

Dernov V., 2022a. Nemors’ki peletsypody mospyns’koyi svity (verkhniy bashkyr) Donets’koho baseynu: systematychnyy sklad, paleoekolohiya ta stratyhrafichne znachennya [Non-marine bivalves from the Mospyne Formation (upper Bashkirian) of the Donets Basin: taxonomy, paleoecology, and stratigraphic significance]. Geologičnij žurnal 380, 34–56.

Dernov V., 2022b. Late Bashkirian ammonoids from the Mospyne Formation of the Donets Basin, Ukraine. Fossil Imprint 78, 489–512.

Dernov V.S., 2022c. The first record of Cyclida (Pancrustacea, Multicrustacea) in the Carboniferous of the Donets Basin, Ukraine. Collection of scientific works of the Institute of Geological Sciences of NAS of Ukraine 15, 78–85.

Dernov V.S., 2022d. Pryzhyttyevi poshkodzhennya cherepashok nemors’kykh dvostulkovykh molyuskiv z karbonu Donets’koho baseynu [Shell injuries of the Carboniferous non-marine bivalves from the Donets Basin]. Paleontological Review 54, 56–64.

Dernov V., 2022e. Biostratyhrafiya bashkyrs’koho yarusu (pensyl’vaniy) Donets’koho baseynu za faunoyu amonoidey [Ammonoid biostratigraphy of the Bashkirian (Pennsylvanian) in the Donets Basin]. Collection of scientific works of the Institute of Geological Sciences of the NAS of Ukraine 15, 123–137.

Dernov V., 2023a. The first record of the trace fossils Cochlichnus from the Pennsylvanian continental and marine deposits in the Donets Basin, Ukraine. Geo&Bio 25, 109–120.

Dernov V., 2023b. Horseshoe crab trace fossils Arborichnus Romano et Meléndez, 1985 from the Bashkirian (Carboniferous) of the Donets Basin, Ukraine. Fossil Imprint 79, 9–25.

Dernov V., 2025a. New records of Pennsylvanian phyllocarid crustaceans (Phyllocarida, Archaeostraca) from the Donets Basin, eastern Ukraine. Geologos 31, 1–16.

Dernov V.S., 2025b. A new record of the myriapod genus Arthropleura Jordan in Jordan et von Meyer, 1854 from the Pennsylvanian of Ukraine. Geologičnij žurnal 390, 33–43.

Dernov V.S. & Poletaev V.I., 2024. New geological and palaeontological data of the Dyakove Group (Carboniferous) and age-related rock formations of the central Donets Basin, Ukraine. Geologičnij žurnal 386, 3–21.

Dernov V.S. & Udovychenko M.I., 2019. K paleobotanicheskoy kharakteristike mospinskoy svity (sredniy karbon, Donbass) [On the palaeobotanical characteristic of the Mospino Formation]. Visnyk of V.N. Karazin Kharkiv National University, Geology, Geography, Ecology 51, 67–82.

Dernov V.S. & Udovychenko N.I., 2021. Nakhodka drevneyshego konkhostraka (Crustacea: Branchiopoda) v karbone Donbassa [A record of the earliest conchostracan crustacean (Crustacea: Branchiopoda) in the Carboniferous of the Donets Basin]. Natural Sciences: Projects, Research, Prospects. Proceedings of the 2nd International Scientific and Practical Conference (Starobilsk, 21–22 December 2021). Starobilsk, 31–33.

Dewar W., 1939. Anthraconauta (Anthracomya) minima (Auctorum) and its associates in the Lancashire coalfield. Bulletin of Geological Survey of Great Britain 1, 47–66.

Dix E. & Pringle J., 1929. On the fossil Xiphosura from the South Wales Coalfield with a note on the myriapod Euphoberia. Summary of Progress, Geological Survey of Great Britain 1928, 90–113.

Dix E. & Pringle J., 1930. Some Coal Measure arthropods from the South Wales Coalfield. Annals and Magazine of Natural History 10, 136–144.

Dunaeva N.M., 1969. Vidkrytyi Donbas [Open Donets Basin]. [In:] Bondarchuk, V.G. (Ed.), Stratygraphiya URSR. Tom 5. Karbon [Stratigraphy of the Ukrainian SSR. Vol. 5. Carboniferous]. Naukova Dumka, Kyiv, 21–48.

Dunn D.L., 1966. New Pennsylvanian conodonts from southwestern United States. Journal of Paleontology 40, 1294–1303.

Eagar R.M.C., 1981. Seasonal sedimentation in the Upper Carboniferous of England: A discussion. The Journal of Geology 89, 765–766.

Eller E.R., 1938. A review of the Xiphosuran genus Belinurus, with the description of a new species, B. alleganyensis. Annals of the Carnegie Museum 27, 129–150.

Eros J.M., Montañez I.P., Osleger D.A., Nemyrovska T.I., Poletaev V.I., Davydov V.I. & Zhykalyak M.V., 2012. Sequence stratigraphy and onlap history of the Donets Basin, Ukraine: insight into Carboniferous icehouse dynamics. Palaeogeography, Palaeoclimatology, Palaeoecology 313–314, 1–25.

Feofilova A.P. & Levenstein M.L., 1963. Osobennosti osadko- i uglenakopleniya v nizhnem i srednem karbone Donetskogo basseyna [Features of sedimentation and coal accumulation in the Lower and Middle Carboniferous of the Donets Basin]. Publishing House of the Academy of Sciences of the USSR, Moscow, 180 pp.

Filipiak P. & Krawczyński W., 1996. Westphalian xiphosurans (Chelicerata) from the Upper Silesia Coal Basin of Sosnowiec, Poland, Acta Palaeontologica Polonica 41, 413–425.

Fisher D.C., 1977. Functional significance of spines in the Pennsylvanian horseshoe crab Euproops danae. Paleobiology 3, 175–195.

Fisher D.C., 1979. Evidence for subaerial activity of Euproops danae (Merostomata, Xiphosurida). [In:] Nitecki M.H. (Ed.), Mazon Creek fossils. Academic Press, New York, 379–447.

Fissunenko O.P., 1987. Srednekamennougol’nyye landshafty Donetskogo basseyna [Middle Carboniferous landscapes of the Donets Basin]. [In:] Bogdanova T.N. & Khozatsky L.I. (Eds.), Palaeontology and geologic history of the palaeobasins Nedra, Leningrad, 92–99.

Fissunenko O.P., 1991. Zonal’naya fitostratigraficheskaya shkala nizhnego i srednego karbona Donetskogo basseyna [Zonal phytostratigraphic scheme of the Lower and Middle Carboniferous of the Donets Basin]. Geologichnij žurnal 3, 55–64.

Harris R.W. & Hollingsworth R.V., 1933. New Pennsylvanian conodonts from Oklahoma. American Journal of Science, Series 5 25, 193–204.

Haug C. Van Roy P., Leipner A., Funch P., Rudkin D.M., Schöllmann L. & Haug J.T., 2012. A holomorph approach to xiphosuran evolution – a case study on the ontogeny of Euproops. Developmental Genes and Evolution 222, 253–268.

Higgins A.C. & Bouckaert J., 1968. Conodont stratigraphy and paleontology of the Namurian of Belgium. Brussels: Service Géologique de Belgique, Mémoires pour servir à l’explication des cartes géologique et miniéres de la Belgique 10, 1–64.

Hind W., 1894–1896. A monograph on Carbonicola, Anthracomya, and Naiadites. Palæontographical Society, London, 182 pp.

Hitchcock E., 1858. Ichnology of New England; a report on the sandstone of the Connecticut Valley, especially its footmarks. W. White, Boston, 220 pp.

Igo H. & Koike T., 1964. Carboniferous conodonts from the Omi limestone, Niigata Prefecture, central Japan (Studies of Asian conodonts, Part I). Transactions and Proceedings of the Palaeontological Society of Japan, New Series 53, 179–193.

Izart A., Briand C., Vaslet D., Vachard D., Coquel R. & Maslo A., 1996. Stratigraphy and sequence stratigraphy of the Moscovian in the Donets Basin. Tectonophysics 268, 189–209.

Jones T.R., 1891. On some Estheriæ and Estheria-like shells from the Carboniferous shales of Western Scotland. Transactions of the Geological Society of Glasgow 9, 79–85.

Jones T.R. & Woodward H., 1899. Contributions to fossil Crustacea. Geological Magazine 6, 388–395.

Karlov N.N., 1948. Prestwiachinella tschernyschevi sp. nov. – novyy predstavnyk Xiphosura z produktyvnoyi tovshchi donetsʹkoho karbonu [Prestwiachinella tschernyschevi sp. nov., a new representative of Xiphosura from the Carboniferous coal-bearing succession of the Donets Basin]. Collection of papers on palaeontology and stratigraphy. Publishing House of the Academy of Sciences of the Ukrainian SSR; Kyiv, 41–47.

Kozitskaya R.I., Kossenko Z.A., Lipnyagov O.M. & Nemirovskaya T.I., 1978. Kamennougol’nyye konodonty Donetskogo basseina [Carboniferous conodonts of the Donets Basin]. Naukova Dumka, Kyiv, 122 pp.

Lamsdell J.C., 2019. Evolutionary history of the dynamic horseshoe crab. [In:] Sitters H.P., Clark J.A., Smith P.A. & Niles L.J. (Eds), Conservation and research on shorebirds and horseshoe crabs in Delaware Bay 1997–2018. International Wader Studies 21, 1–14.

Lamsdell J.C., 2020. The phylogeny and systematics of Xiphosura. PeerJ 8: e10431.

Lamsdell J.C., 2022. One name to rule them all: Belinurus trilobitoides (Buckland, 1837) is senior synonym to fourteen named species. Journal of Paleontology 96, 237–241.

Lamsdell J.C. & Clapham M.E., 2021. Belinurus Bronn, 1839 (Chelicerata, Xiphosura) has priority over Bellinurus Pictet, 1846. Journal of Paleontology 95, 1352–1355.

Latreille P.A., 1802. Histoire naturelle générale et particulière des crustacés et des insectes: ouvrage faisant suite aux Oeuvres de Leclerc de Buffon, et partie du Cours complet d’histoire naturelle rédigé par C.S. Sonnini, 7. F. Dufart, Paris, 413 pp.

Linnaeus C., 1758. Systema Naturae. Vol. 1. 10th edition. Laurentius Salvius, Holmiae, 824 pp.

Leach W., 1819. Entomostracés. [In:] Levrault F. (Ed.), Dictionnaires des Sciences Naturelles, 14. Strasbourg, pp. 524–543.

Logvinenko N.V., 1953. Litologiya i paleogeografiya produktivnoy tolshchi donetskogo karbona [Lithology and palaeogeography of the Carboniferous coal-bearing deposits of the Donets Basin]. Publishing House of the Kharkiv University, Kharkiv, 436 pp.

Lucas S.G., Lerner A.J., Dimichele W.A., Cantrell A.K., Suazo T.L. & Chaney D.S., 2014. Xiphosurid fossils from the Pennsylvanian Beeman Formation, Otero County, New Mexico. New Mexico Geological Society Guidebook, 65th Field Conference, Geology of the Sacramento Mountains Region, 311–314.

Lustri L., Laibl L. & Bicknell R.D.C., 2020. A revision of Prolimulus woodwardi Fritsch, 1899 with comparison to other highly paedomorphic belinurids. PeerJ 9, e10980.

Meek E.B., 1867. Note on a new genus of fossil Crustacea. The American Journal of Science and Arts (Series 2) 43, 394–395.

Meek F.B. & Worthen A.H., 1865. Notice of some new types of organic remains, from the Coal Measures of Illinois. Proceedings of the Academy of Natural Sciences 12, 41–45.

Mikulic D.G., 1998. Xiphosura. [In:] Shabica C.W. & Hays A.A. (Eds.), Richardson’s guide to the fossil fauna of Mazon Creek. Northeastern Illinois University Press, Chicago, 134–139.

Müller O., 1785. Entomostraca, seu, Insecta testacea quae in aquis Daniae et Norvegie reperit, descripsit et iconibus illustravit. F.W. Thiele, Hauniae, 135 pp.

Mychko E.V. & Dernov V.S., 2019. Novyye nakhodki trilobitov v srednem karbone Donetskogo basseyna [New record of trilobites in the middle Carboniferous of the Donets Basin]. Bulletin of the Moscow Society of Naturalists (Geological Section) 94, 52–63.

Nemyrovska T.I., 1999. Bashkirian conodonts of the Donets Basin, Ukraine. Scripta Geologica 119, 1–116.

Nemyrovska T.I., 2017. Late Mississippian–Middle Pennsylvanian conodont zonation of Ukraine. Stratigraphy 14, 299–318.

Nemyrovska T.I., & Yefimenko V.I., 2013. Seredniy karbon (nyzniy pensyl’vaniy) [Middle Carboniferous (Lower Pennsylvanian). [In:] Gozhyk P.F. (Ed.), Stratyhrafiya verkhnʹoho proterozoyu ta fanerozoyu Ukrayiny. Tom 1. Stratyhrafiya verkhnʹoho proterozoyu, paleozoyu ta mezozoyu Ukrayiny [Stratigraphy of the Upper Proterozoic and Phanerozoic of Ukraine. Vol. 1. Stratigraphy of the Upper Proterozoic, Palaeozoic and Mesozoic]. LAT&K, Kyiv, 283–303.

Newberry J.S., 1856. Description of several new genera and species of fossil fishes, from the Carboniferous strata of Ohio. Proceedings of the Academy of Natural Sciences of Philadelphia 8, 96–100.

Novik E.O., 1974. Zakonomernosti razvitiya kamennougol’noy flory yuga Yevropeyskoy chasti SSSR [Development of the Carboniferous flora of the south part of the European part of the USSR]. Naukova Dumka, Kyiv, 139 pp.

Packard A.S., 1886. On the Carboniferous xiphosurous fauna of North America. Memoirs of the National Academy of Sciences 3, 143–157.

Pictet F.J., 1854. Traite élémentaire de paléontologie. Vol. 2. Bailliere, Paris, 727 pp.

Poletaev V.I. & Vdovenko M.V., 2013. Nyzhniy karbon (Missisipiy) [Lower Carboniferous (Mississippian)]. [In:] Gozhik P.F. (Ed.), Stratygraphiya verkhnyogo proterozoyu i fanerozoyu Ukrainy. Tom 1. Stratygraphiya verkhnyogo proterozoyu, paleozoyu i mezozoyu [Stratigraphy of the Upper Proterozoic and Phanerozoic of Ukraine. Volume 1. Stratigraphy of the Upper Proterozoic, Palaeozoic and Mesozoic]. LAT&K, Kyiv, 250–283.

Poletaev V.I., Vdovenko M.V., Shchoholev O.K., Boyarina N.I., & Makarov I.A., 2011. Stratotypy rehional’nykh stratyhrafichnykh pidrozdiliv karbonu i nyzhn’oyi permi Dono-Dniprovs’koho prohynu [Stratotypes of the Carboniferous and Lower Permian regional stratigraphic units of the Don-Dnipro Downwarp]. Logos, Kyiv, 234 pp.

Popov A.V., 1979. Kamennougol’nyye ammonoidei Donbassa i ikh stratigraficheskoye znacheniye [Carboniferous ammonoids of the Donets Basin and their stratigraphic significance]. Transactions of the All-Union Geological Institute, New series 220, 1–119.

Posudiyevsky A.B., 1977. K izucheniyu fatsial’nogo sostava porod neposredstvennoy krovli ugol'nykh plastov v Donbasse [On the study of depositional environments of the roof shales of the coal beds in the Donets Basin]. Lithology and mineral resources 3, 156–159.

Prantl F. & Přibyl A., 1955. Ostrorepi (Xiphosura) ceskoslovenskeho karbonu. Sborník Ústředního ústavu geologického 22, 379–424.

Pruvost P., 1930. La faune continentale du terrain houiller de la Belgique. Mémoires du Musée royal d'histoire naturelle de Belgique 44, 103–282.

Reznikov A.I., 1993. D’yakovskaya seriya i yeye polozheniye v stratigraficheskoy skheme Donbassa [The Dyakove Group and its stratigraphic position in the Carboniferous succession of the Donets Basin]. Geologìčnij žurnal 262, 52–57.

Roemer F., 1883. Über eine Art der limuliden-gattung Belinurus aus dem Steinkohlengebirge Oberschlesiens. Zeitschrift der Deutschen Geologischen Gesellschaft 35, 429–432.

Sachsenhofer R.F., Privalov V.A., & Panova E.A., 2012. Basin evolution and coal geology of the Donets Basin (Ukraine, Russia): An overview. International Journal of Coal Geology 89, 26–40.

Schultka S., 1994. Bellinurus cf. truernanii (Merostomata) aus dem tiefen Oberkarbon (Namur B/C) von Frodenberg (Nordrhein-Westfalen, Deutschland). Paläontologische Zeitschrift 68, 339–349.

Schwarzbach M., 1962. Die Merostomata aus dem niederrheinisch-westfälischen Oberkarbon. Das Karbon der subvariscischen Saumsenke. Ein Symposium. Teil 2. Fortschritte in der Geologie von Rheinland und Westfalen 3, 803–818.

Selden P.A., Simonetto L. & Marsiglio G., 2019. An effaced horseshoe crab (Arthropoda: Chelicerata: Xiphosura) from the Upper Carboniferous of the Carnic Alps (Friuli, NE Italy). Rivista Italiana di Paleontologia e Stratigrafia 125, 333–342.

Shpinev E.S., 2018. New data on Carboniferous xiphosurans (Xiphosura, Chelicerata) of the Donets Coal Basin. Paleontological Journal 52, 271–283.

Shulga V.F., Zdanowski A., Zaitseva L.B., Ivanova A.V., Ivanina A.V., Korol N.D., Kotasova A., Kotas A., Kostyk I.E., Lelyk B.I., Miger T., Manichev V.I., Matrofailo M.N., Ptak B., Savchuk V.S., Sedaeva G.M. & Stepanenko Ya.G., 2007. Korrelyatsiya karbonovykh uglenosnykh formatsiy L'vovsko-Volynskogo i Lyublinskogo basseynov [Correlation of the Carboniferous coal-bearing formations of the Lviv-Volyn and Lublin basins]. Varta, Kyiv, 427 pp.

Siegfried P., 1972. Ein Schwertschwanz (Merostomata, Xiphosurida) aus dem Oberkarbon von Ibbenbüren/Westf. Paläontologische Zeitschrift 46, 180–185.

Stainier X., 1938. Belinurus et Vetacapsula nouveaux du houiller Belge. Bulletin du Musée royal d'Histoire naturelle de Belgique 14, 1–13.

Tashman J.N., Feldmann R.M. & Schweitzer C.E., 2019. Morphological variation in the Pennsylvanian horseshoe crab Euproops danae (Meek & Worthen, 1865) (Xiphosurida, Euproopidae) from the lower Mercer Shale, Windber, Pennsylvania, USA. Journal of Crustacean Biology 39, 396–406.

Trueman A.E. & Weir J., 1948. A monograph of British Carboniferous non-marine Lamellibranchia. Part IV. Monographs of the Palaeontographical Society 102, 75–100.

Ustinovsky Yu.B., 1993. Kayal’skiy etap (pozdniy bashkir) [Kayalian stage (Late Bashkirian)]. [In:] Tsegelnyuk P.D. (Ed.), Geologicheskaya istoriya territorii Ukrainy [Geological history of Ukraine]. Naukova Dumka, Kyiv, 141–144.

Van Hinsbergen D.J.J., Abels H.A., Bosch W., Boekhout F., Kitchka A., Hamers M., van der Meer D.G, Geluk M. & Stephenson R.A., 2015. Sedimentary geology of the middle Carboniferous of the Donbas region (Dniepr-Donets basin, Ukraine). Scientific Reports 5 (9099).

Warth M., 1967. Die nichtmarinen Muscheln des Westfal A unter besonderer Berücksichtigung des Ruhrkarbons und einige grundlegende Erkenntnisse zur Taxonomie. Westdeutscherverlag, Kӧln, 125 s.

Weber V.N., 1933. Trilobity Donetskogo basseyna [Trilobites of the Donets Basin]. Transactions of the All-Union Geological Exploration Department of NKTP of the USSR 225, 1–95.

Wignall P.B. & Sims M.J., 1990. Pseudoplankton. Palaeontology 33, 359–378.

Williamson W.C., 1849. On the microscopic structure of the scales and dermal teeth of some ganoid and placoid fish. Philosophical Transactions of the Royal Society of London 139, 435–474.

Woodward H., 1867. On some points in the structure of the Xiphosura having reference to their relationship with the Eurypteridae. Quarterly Journal of the Geological Society of London 23, 28–37.

Woodward H., 1893. Note on a new British species of Cyclus from the Coal Measures of Bacup, Lancashire. Geological Magazine 10, 28–29.

Woodward H., 1918. Fossil arthropods from the Carboniferous rocks of Cape Breton, Nova Scotia; and from the Upper Coal Measures, Sunderland, England. Geological Magazine 5, 462–471.

Woodward H. & Etheridge R., 1873. Explanation of sheet 23. Memoirs of the Geologic Survey, Scotland, 1–99.

Wright W.B., 1934. The fresh-water fauna of the Lower Measures of Lancashire. Summary of progress of the Geological Survey of Great Britain and the Museum of Practical Geology for the year 1933 2, 8–23.

Yefimov I.N., 1934. Raspredeleniye organicheskikh ostatkov v krovle plastov kamennykh ugley Donetskogo basseyna [Distribution of fossils in the roof shales of the coal beds in the Donets Basin]. Reports of the Academy of Sciences of the USSR 6, 123–125.

Zalessky M.D., 1907. Materialy po kamennougol'noy flore Donetskogo basseyna. I. Rastitel'nyye ostatki kollektsii V. Domgera [Materials on the Carboniferous flora of the Donets Basin. I. Plant remains of V. Domher’s collection]. Proceedings of the Geological Committee 26, 351–422.

Zaritskiy P.V., 1970. Mineralogiya i geokhimiya diageneza uglenosnykh otlozheniy (na primere Donetskogo basseyna). Chast’ 1. [Mineralogy and geochemistry of diagenesis of coal-bearing deposits (based on materials from the Donets Basin). Part 1]. Kharkiv University Press, Kharkiv, 225 pp.

Zhemchuzhnikov Yu.A., Yablokov V.S., Bogolyubova L.I., Botvinkina L.N., Feofilova A.P., Ritenberg M.I., Timofeev P.P. & Timofeeva Z.V., 1960. Stroyeniye i usloviya nakopleniya glavnykh uglenosnykh svit i ugol'nykh plastov srednego karbona v Donetskom basseyne. Chast’ 2 [Structure and depositional conditions of the main coal-bearing formations and coal seams of the middle Carboniferous in the Donets Basin. Part 2]. Transactions of the Geological Institute 15, 1–247.

Zittel K.A. von & Eastman C.R., 1913. Textbook of Palaeontology. Macmillan, London, 839 pp.