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The first occurrence of the earliest species of Acadoparadoxides outside West Gondwana (Cambrian; Holy Cross Mountains, Poland)



Two hundred and eighty specimens of paradoxidids from two localities in the Holy Cross Mountains (Poland) have been reanalysed using morphometric techniques. Revision of the dataset provided evidence for the presence of two endemic taxa: Acadoparadoxides kozlowskii (Orłowski, 1959) and Acadoparadoxides samsonowiczi (Orłowski, 1959), both belonging to the earliest group of Acadoparadoxides, initially considered to be present only in Gondwanan successions. Thus, this is the first description of the members of this group outside West Gondwana. The taxonomic revision, and the presence of representatives of the earliest acadoparadoxidines coupled with the absence of taxa typical of Scandinavia that were previously described from this locality have resulted in the modification of the biostratigraphic scheme hitherto used in the Holy Cross Mountains. The newly established A. samsonowiczi – A. kozlowskii Assemblage Zone is correlated with the Morocconus notabilis Zone of Morocco, and thus represents the Cambrian Series 2 and 3 boundary interval. Links with West Gondwana challenge the existing palaeogeographic interpretations for the southern part of the Holy Cross Mountains and point to an urgent need to revise the position of the Małopolska Block during the middle Cambrian.


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Adams, D. C., Collyer, M. L., Kaliontzopoulou, A. & Sherratt, E. 2016. Geomorph: Software for Geometric Morphometric Analyses. R Package Version 3.0.2. Available at (accessed 25 August 2016).
Adams, D. C. & Otárola-Castillo, E. 2013. geomorph: an R package for the collection and analysis of geometric morphometric shape data. Methods in Ecology and Evolution 4, 393–9.
Álvaro, J. J., Esteve, J. & Zamora, S. 2017. Morphological assessment of the earliest paradoxidid trilobites (Cambrian Series 3) from Morocco and Spain. Geological Magazine 154, 1306–28.
Angielczyk, K. D. & Sheets, H. D. 2007. Investigation of simulated tectonic deformation in fossils using geometric morphometrics. Paleobiology 33, 125–48.
Asklund, B. & Thorslund, P. 1935. Fjällkedjerandens bergbyggnad i norra Jämtland och Ångermanland. Sveriges Geologiska Undersökning C 382, 1110.
Barrande, J. 1846. Notice préliminaire sur se système silurien et les trilobites de Bohême. Prague: C. L. Hirschfeld, 97 pp.
Barrande, J. 1852. Système silurien du centre de la Bohême. 1ère partie: Recherches paléontologiques. Vol. I. Crustacés: Trilobites. Prague: issued by the author, 935 pp.
Bassett, M. G., Popov, L. E. & Holmer, L. E. 2002. Brachiopods: Cambrian–Tremadoc precursors to Ordovician radiation events. In Palaeobiogeography and Biodiversity Change: The Ordovician and Mesozoic–Cenozoic Radiations (eds Crame, J. A. & Owen, A. W.), pp. 1323. Geological Society of London, Special Publication no. 194.
Bednarczyk, W. 1970. Trilobites fauna of the Lower Paradoxides oelandicus stage from the Brzechów area in the western part of the Świętokrzyskie Mts. Bulletin de l'Académie Polonaise des Sciences, Série des sciences chimiques, géologiques et géographiques 18, 2935.
Bednarczyk, W., Lendzion, K. & Orłowski, S. 1990. Rodzina Paradoxididae Hawle et Corda, 1847. In Budowa Geologiczna Polski. Tom III. Atlas Skamieniałości Przewodnich i Charakterystycznych. Część 1a. Paleozoik Starszy (z Proterozoikiem Górnym). Ryfej, Wend, Kambr, pp. 58–61. Warsaw, Poland: Państwowy Instytut Geologiczny.
Belka, Z., Valverde-Vaquero, P., Dörr, W., Ahrendt, H., Wemmer, K., Franke, W. & Schäfer, J. 2002. Accretion of first Gondwana-derived terranes at the margin of Baltica. Palaeozoic Amalgamation of Central Europe 201, 1936.
Clarke, K. R. 1993. Non-parametric multivariate analyses of changes in community structure. Australian Journal of Ecology 18, 117–43.
Czarnocki, J. 1927a. Kambr i jego fauna w środkowej części Gór Świętokrzyskich. Sprawozdania Państwowego Instytutu Geologicznego 4, 189208.
Czarnocki, J. 1927b. Le Cambrien et la faune cambrienne de la partie moyenne du Massif de Swiety Krzyz (Ste Croix). In Extrait du Compte-Rendu XIVème Congrès Géologique International, pp. 1–18.
Dean, W. T. 2005. Trilobites from the Çal Tepe Formation (Cambrian), near Seydişehir, Central Taurides, Southwestern Turkey. Turkish Journal of Earth Sciences 14, 171.
Dean, W. T. & Rushton, A. W. A. 1997. Superfamily Paradoxidoidea. In Treatise on Invertebrate Paleontology, Part O, Arthropoda 1. Trilobita, Revised (eds Whittington, H. B., Chatterton, B. D. E., Speyer, S. E., Fortey, R. A., Owens, R. M., Chang, W. T., Dean, W. T., Geyer, G., Jell, P. A., Laurie, J. R., Palmer, A. R., Repina, L. N., Rushton, A. W. A., Shergold, J. H., Clarkson, E. N. K., Wilmot, N. V. & Kelly, S. R. A.), pp. O470–81. Lawrence, KS: University of Kansas Press / Boulder, CO: Geological Society of America.
Drygant, D. & Konstantinenko, L. 2001. A new trilobite genus from the Cambrian of the Carpathian Foredeep. Paleontological Collection 33, 2831 (in Ukrainian).
Dzik, J. & Orłowski, S. 1995. Primitive ctenocystoid echinoderm from the earliest Middle Cambrian of Poland. Annales de Paléontologie 81, 1735.
Elicki, O. & Geyer, G. 2013. The Cambrian trilobites of Jordan – taxonomy, systematic and stratigraphic significance. Acta Geologica Polonica 63, 156.
Esteve, J. 2014. Intraspecific variability in paradoxidid trilobites from the Purujosa trilobite assemblage (middle Cambrian, northeast Spain). Acta Palaeontologica Polonica 59, 215–40.
Geyer, G. 1993. The giant Cambrian trilobites of Morocco. Beringeria 8, 71107.
Geyer, G. 1998. Intercontinental, trilobite-based correlation of the Moroccan early Middle Cambrian. Canadian Journal of Earth Sciences 35, 374401.
Geyer, G. 2016. Taxonomy of the ‘Micmacca’ group, new Cambrian Chengkouiidae (Trilobita) from Morocco, and their bearing on intercontinental correlation. Australasian Palaeontological Memoirs 49, 329–93.
Geyer, G. & Landing, E. 2001. Middle Cambrian of Avalonian Massachusetts: stratigraphy and correlation of the Braintree trilobites. Journal of Paleontology 75, 116–35.
Geyer, G. & Landing, E. 2004. A unified Lower–Middle Cambrian chronostratigraphy for West Gondwana. Acta Geologica Polonica 54, 179218.
Geyer, G. & Landing, E. 2006. Latest Ediacaran and Cambrian of the Moroccan Atlas regions. In Ediacaran–Cambrian Depositional Environments and Stratigraphy of the Western Atlas Regions (eds Geyer, G. & Landing, E.), pp. 746. University College London Maghreb Petroleum Research Group, Infracambrian / Early Palaeozic Field Guide Series 1 and Beringeria Special Issue 6.
Geyer, G. & Vincent, T. 2015. The Paradoxides puzzle resolved: the appearance of the oldest paradoxidines and its bearing on the Cambrian Series 3 lower boundary. Paläontologische Zeitschrift 89, 335–98.
Gozalo, R., Dies Alvarez, M. E., Gamez Vintaned, J. A., Zhuravlev, A., Bauluz, B., Subías, I., Chirivella Martorell, J. B., Mayoral, E., Gursky, H. J., Andrés, J. A. & Liñán, E. 2013. Proposal of a reference section and point for the Cambrian Series 2–3 boundary in the Mediterranean subprovince in Murero (NE Spain) and its intercontinental correlation. Geological Journal 48, 142–55.
Green, J. 1834. Descriptions of some new North American trilobites. American Journal of Science 25, 334–7.
Haber, A. 2011. OSymm. Available at (accessed 29 April 2015).
Hawle, I. & Corda, A. J. C. 1847. Prodrom einer Monographie der böhmischen Trilobiten. Prague: J. G. Calve, 176 pp.
Hughes, N. C. & Jell, P. A. 1992. A statistical/computer-graphic technique for assessing variation in tectonically deformed fossils and its application to Cambrian trilobites from Kashmir. Lethaia 25, 317–30.
Hupé, P. 1953. Contribution a l’étude du Cambrien inférieur et du Précambrien III de l'Anti-Atlas marocain. Notes et Mémoires de la Service géologique du Maroc 103, 1402.
Jendryka-Fuglewicz, B. 1992. Analiza porównawcza ramienionogów z utworów kambru Gór Świętokrzyskich i platformy prekambryjskiej w Polsce [Comparison analysis of brachiopods from the Cambrian deposits of the Holy Cross Mountains]. Przegląd Geologiczny 40, 150–5.
Kerner, A. & Debrenne, F. 2013. The role of Archaeocyatha in Cambrian biostratigraphy and biogeography. In Early Palaeozoic Biogeography and Palaeogeography (eds Harper, D.A.T. & Servais, T.), pp. 5966. Geological Society of London, Memoir 38.
Klingenberg, C. P., Barluenga, M. & Meyer, A. 2002. Shape analysis of symmetric structures: quantifying variation among individuals and asymmetry. Evolution; International Journal of Organic Evolution 56, 1909–20.
Kowalczewski, Z., Żylińska, A. & Szczepanik, Z. 2006. Kambr w Górach Świętokrzyskich. In LXXVII Zjazd Naukowy Polskiego Towarzystwa Geologicznego, Ameliówka k. Kielc 28–30 czerwca 2006 r., pp. 14–27. Warsaw, Poland: Państwowy Instytut Geologiczny.
Laibl, L., Esteve, J. & Fatka, O. 2017. Giant postembryonic stages of Hydrocephalus and Eccaparadoxides and the origin of lecithotrophy in Cambrian trilobites. Palaeogeography, Palaeoclimatology, Palaeoecology 470, 109–15.
Landing, E., Geyer, G., Brasier, M. D. & Bowring, S. A. 2013. Cambrian evolutionary radiation: context, correlation, and chronostratigraphy – overcoming deficiencies of the first appearance datum (FAD) concept. Earth-Science Reviews 123, 133–72.
Masiak, M. & Żylińska, A. 1994. Burgess Shale-type fossils in Cambrian sandstones of the Holy Cross Mountains. Acta Palaeontologica Polonica 39, 329–40.
Matthew, G. F. 1887. Illustrations of the fauna of the St. John Group, No. IV. On the smaller-eyed trilobites of Division I, with a few remarks on the species of the higher divisions of the group. Canadian Record of Science 2, 357–63.
Meyer, D., Dimitriadou, E., Hornik, K., Weingessel, A. & Leisch, F. 2017. e1071: Misc Functions of the Department of Statistics, Probability Theory Group (Formerly: E1071), TU Wien. Available at (accessed 5 December 2017).
Narkiewicz, M. & Petecki, Z. 2017. Basement structure of the Paleozoic Platform in Poland. Geological Quarterly 61, 502–20.
Nawrocki, J., Dunlap, J., Pecskay, Z., Krzemiński, L., Żylińska, A., Fanning, M., Kozłowski, W., Salwa, S., Szczepanik, Z. & Trela, W. 2007. Late Neoproterozoic to Early Paleozoic paleogeography of the Holy Cross Mountains (Central Europe): an integrated approach. Journal of the Geological Society of London 164, 405–23.
Nielsen, A. T. & Schovsbo, N. H. 2015. The regressive Early-Mid Cambrian ‘Hawke Bay Event’ in Baltoscandia: epeirogenic uplift in concert with eustasy. Earth-Science Reviews 151, 288350.
Nowicki, J. 2015. morphoutils. Available at
Nowicki, J. 2016. geomorphotest. Available at
Orłowski, S. 1957. On the presence of Paradoxides ölandicus beds in the Holy Cross Mountains. Bulletin de l'Académie Polonaise des Sciences, Série des sciences chimiques, géologiques et géographiques 5, 769–72.
Orłowski, S. 1959a. Paradoxidae from lower Middle Cambrian strata in the vicinity of Sandomierz (Central Poland). Bulletin de l'Académie Polonaise des Sciences, Série des sciences chimiques, géologiques et géographiques 7, 441–6.
Orłowski, S. 1959b. Ellipsocephalidae from the Lower Beds of the Middle Cambrian in the vicinity of Sandomierz (Central Poland). Bulletin de l'Académie Polonaise des Sciences, Série des sciences chimiques, géologiques et géographiques 7, 515–20.
Orłowski, S. 1964. Kambr środkowy i jego fauna we wschodniej części Gór Świętokrzyskich [Middle Cambrian and its fauna in the eastern part of the Holy Cross Mts.]. Studia Geologica Polonica 16, 194.
Orłowski, S. 1975. Jednostki litostratygraficzne kambru i górnego prekambru w Górach Świetokrzyskich [Cambrian and Upper Precambrian lithostratigraphic units in the Holy Cross Mts.] Acta Geologica Polonica 25, 431–48.
Orłowski, S. 1985. New data on the Middle Cambrian trilobites and stratigraphy in the Holy Cross Mts. Acta Geologica Polonica 35, 251–63.
Orłowski, S. 1992. Cambrian stratigraphy and stage subdivision in the Holy Cross Mountains, Poland. Geological Magazine 129, 471–4.
R Core Team. 2016. R: A Language and Environment for Statistical Computing. Available at
Reed, F. R. C. 1910. The Cambrian fossils of Spiti. Palaeontologia Indica Series 15 7, 170.
Richter, R. 1932. Crustacea (Paläontologie). In Handwörterbuch der Naturwissenschaften, 2. Auflage 2. (eds Dittler, R., Joos, D., Korschelt, E., Linck, G., Oltmanns, F. & Schaum, K.), pp. 840–64. Jena: Fischer.
Rohlf, F. J. 2016. TPS Software Series. Available at (accessed 13 April 2015).
Rushton, A. W. A. & Weidner, T. 2007. The Middle Cambrian paradoxidid trilobite Hydrocephalus from Jämtland, central Sweden. Acta Geologica Polonica 57, 391401.
Rushton, A. W. A., Weidner, T. & Ebbestad, J. O. R. 2016. Paradoxidid trilobites from a mid-Cambrian (Series 3, stage 5) limestone concretion from Jämtland, central Sweden. Bulletin of Geosciences 91, 515–52.
Samsonowicz, J. 1959. On Strenuaeva from Lower Cambrian in Klimontów Anticlinorium. Bulletin de l'Académie Polonaise des Sciences, Série des Sciences Chimiques, Géologiques et Géographiques 7, 521–4.
Sdzuy, K. 1958. Neue Trilobiten aus dem Mittelkambrium von Spanien. Senckenbergiana lethaea 39, 235–53.
Sdzuy, K. 1967. Trilobites del Cámbrico Medio de Asturias. Trabajos de Geología, Faculdad de Ciencias, Universidad de Oviedo 1, 77133.
Servais, T., Danelian, T., Alexander, D., Harper, T. & Munnecke, A. 2014. Possible oceanic circulation patterns, surface water currents and upwelling zones in the Early Palaeozoic. GFF 136, 229–33.
Sjögren, A. 1872. Om några försteningar i Ölands kambriska lager. Geologiska Föreningen i Stockholm Förhandlingar 1 (5), 6780.
Smith, A. G. 2001. Paleomagnetically and tectonically based global maps for Vendian to Mid-Ordovician Time. In The Ecology of the Cambrian Radiation, pp. 1146. NewYork: Columbia University Press.
Šnajdr, M. 1957. O nových trilobitech z českého kambria. Věstnik Ústředniho ústavu geologického 32, 235–44.
Šnajdr, M. 1958. Trilobiti českého středního kambria. Rozpravy Ústředniho ústavu geologického 24, 1280.
Šnajdr, M. 1986. Two new paradoxid trilobites from Jince Formation (Middle Cambrian, Czechoslovakia). Věstnik Ústředniho ústavu geologického 61, 169–74.
Šnajdr, M. 1987. The genera Paradoxides Brongniart and Hydrocephalus Barrande (Trilobita). Věstnik Ústředniho ústavu geologického 62, 97104.
Solov'ev, A. I. 1969. Novye vidy Paradoxides (trilobity) iz goryuchikh slantsev amginskogo yarusa Severnoy Yakutii. Uchenie zapiyski, NIIGA, paleontologiya i biostratigrafiya 25, 920.
Srivastava, D. C. & Shah, J. 2006. Digital method for strain estimation and retrodeformation of bilaterally symmetric fossils. Geology 34, 593–6.
Stasińska, A. 1960. Velumbrella czarnockii n. gen., n. sp. – méduse du Cambrien Inférieur des Monts de Sainte-Croix. Acta Palaeontologica Polonica 5, 337–44.
Sundberg, F. A., Geyer, G., Kruse, P. D., McCollum, L. B., Pegel’, T. V., Żylińska, A. & Zhuravlev, A. Y. 2016. International correlation of the Cambrian Series 2–3, Stages 4–5 boundary interval. Australasian Paleontological Memoirs 49, 83124.
Torsvik, T. H. & Cocks, L. R. M. 2013. New global palaeogeographical reconstructions for the Early Palaeozoic and their generation. In Early Palaeozoic Biogeography and Palaeogeography (eds Harper, D.A.T. & Servais, T.), pp. 524. Geological Society of London, Memoir 38.
Walch, J. E. I. 1771. Die Naturgeschichte der Versteinerungen zur Erläuterung der Knorrischen Sammlung von Merkwürdigkeiten der Natur. Nürnberg: Felßecker, 303 pp.
Walczak, A. & Belka, Z. 2017. Fingerprinting Gondwana versus Baltica provenance: Nd and Sr isotopes in Lower Paleozoic clastic rocks of the Małopolska and Łysogóry terranes, southern Poland. Gondwana Research 45, 138–51.
Weidner, T. R., Rushton, A. W. A. & Ebbestad, J. O. R. 2014. A paradoxidid–agnostoid fauna from the mid-Cambrian (Stage 5) of the Caledonian Lower Allochthon on Tåsjöberget, Ångermanland, Sweden. GFF 136, 513–30.
Westergård, A. H. 1936. Paradoxides oelandicus Beds of Öland. Sveriges Geologiska Undersökning C 394, 166.
Whittington, H. B. 1997. Morphology of the exoskeleton. In Treatise on Invertebrate Paleontology, Part O, Arthropoda 1. Trilobita, Revised (eds. Whittington, H. B., Chatterton, B. D. E., Speyer, S. E., Fortey, R. A., Owens, R. M., Chang, W. T., Dean, W. T., Geyer, G., Jell, P. A., Laurie, J. R., Palmer, A. R., Repina, L. N., Rushton, A. W. A., Shergold, J. H., Clarkson, E. N. K., Wilmot, N. V. & Kelly, S. R. A.), pp. O1–O67. Lawrence, KS: University of Kansas Press / Boulder, CO: Geological Society of America.
Winrow, P. & Sutton, M. D. 2014. Lingulate brachiopods and the Early Palaeozoic history of the Iapetus Ocean. Lethaia 47, 456–68.
Zelditch, M. L., Swiderski, D. L. & Sheets, H. D. 2012. Geometric Morphometrics for Biologists: A Primer. Second edition. London: Elsevier Academic Press, 488 pp.
Żylińska, A. 2013a. Ewolucja zespołów trylobitowych z drugiego i trzeciego oddziału kambru Gór Świętokrzyskich i ich znaczenie biogeograficzne (Evolution of trilobite assemblages from the Cambrian Series 2 and 3 of the Holy Cross Mountains and their biogeographic significance). Przegląd Geologiczny 61, 30–9.
Żylińska, A. 2013b. The oldest Cambrian trilobites from the Holy Cross Mountains, Poland: taxonomic, stratigraphic and biogeographic reappraisal. Acta Geologica Polonica 63, 5787.
Żylińska, A., Kin, A. & Nowicki, J. 2013. Application of morphometric techniques for taxonomic revision of Berabichia oratrix (Orłowski, 1985) (Trilobita, Cambrian) from the Holy Cross Mountains, Poland. Geodiversitas 35, 505–28.
Żylińska, A. & Masiak, M. 2007. Cambrian trilobites from Brzechów, Holy Cross Mountains (Poland) and their significance in stratigraphic correlation and biogeographic reconstructions. Geological Magazine 144, 661–86.
Żylińska, A. & Szczepanik, Z. 2009. Trilobite and acritarch assemblages from the Lower – Middle Cambrian boundary interval in the Holy Cross Mountains (Poland). Acta Geologica Polonica 59, 413–58.



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