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The Giant Pliosaurid That Wasn't—Revising the Marine Reptiles From The giant pliosaurid that wasn’t—revising the marine reptiles from the Kimmeridgian, Upper Jurassic, of Krzyżanowice, Poland DANIEL MADZIA, TOMASZ SZCZYGIELSKI, and ANDRZEJ S. WOLNIEWICZ Madzia, D., Szczygielski, T., and Wolniewicz, A.S. 2021. The giant pliosaurid that wasn’t—revising the marine reptiles from the Kimmeridgian, Upper Jurassic, of Krzyżanowice, Poland. Acta Palaeontologica Polonica 66 (1): 99–129. Marine reptiles from the Upper Jurassic of Central Europe are rare and often fragmentary, which hinders their precise taxonomic identification and their placement in a palaeobiogeographic context. Recent fieldwork in the Kimmeridgian of Krzyżanowice, Poland, a locality known from turtle remains originally discovered in the 1960s, has reportedly provided additional fossils thought to indicate the presence of a more diverse marine reptile assemblage, including giant pliosaurids, plesiosauroids, and thalattosuchians. Based on its taxonomic composition, the marine tetrapod fauna from Krzyżanowice was argued to represent part of the “Matyja-Wierzbowski Line”—a newly proposed palaeobiogeographic belt comprising faunal components transitional between those of the Boreal and Mediterranean marine provinces. Here, we provide a de- tailed re-description of the marine reptile material from Krzyżanowice and reassess its taxonomy. The turtle remains are proposed to represent a “plesiochelyid” thalassochelydian (Craspedochelys? sp.) and the plesiosauroid vertebral centrum likely belongs to a cryptoclidid. However, qualitative assessment and quantitative analysis of the jaws originally referred to the colossal pliosaurid Pliosaurus clearly demonstrate a metriorhynchid thalattosuchian affinity. Furthermore, these me- triorhynchid jaws were likely found at a different, currently indeterminate, locality. A tooth crown previously identified as belonging to the thalattosuchian Machimosaurus is here considered to represent an indeterminate vertebrate. The revised taxonomy of the marine reptiles from Krzyżanowice, as well as the uncertain provenance of the metriorhynchid specimen reported from the locality, cast doubt on the palaeobiogeographic significance of the assemblage. Key words: Pliosauridae, Metriorhynchidae, Crocodylomorpha, Thalassochelydia, Kimmeridgian, Jurassic, Poland. Daniel Madzia [[email protected]; ORCID: https://orcid.org/0000-0003-1228-3573], Tomasz Szczygielski [[email protected]; ORCID: https://orcid.org/0000-0001-5108-8493], and Andrzej S. Wolniewicz [wolnie- [email protected]; ORCID: https://orcid.org/0000-0002-6336-8916], Institute of Paleobiology, Polish Academy of Sciences, ul. Twarda 51/55, 00-818 Warszawa, Poland. Received 21 July 2020, accepted 14 September 2020, available online 12 January 2021. Copyright © 2021 D. Madzia et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Testudinata and Lepidosauria, respectively (Bardet et al. Introduction 2014; Anquetin et al. 2017). The Late Jurassic was an important time interval in the evo- In Europe, abundant and well-preserved fossils of Late lutionary history of several clades of Mesozoic marine rep- Jurassic marine reptiles are known from the Kimmeridge tiles. Some groups, such as ophthalmosaurid ichthyosaurs, Clay Formation (Kimmeridgian) of the United Kingdom plesiosaurs (pliosaurids and cryptoclidids, in particular), (e.g., Owen 1842; Seeley 1869, 1875; Lydekker 1889; And- and thalattosuchian crocodylomorphs had already become rews 1921; Tarlo 1960; Brown 1981; Brown et al. 1986; globally distributed and underwent significant taxonomic Tay lor and Cruickshank 1993; Sassoon et al. 2012; Young and ecomorphological diversification by the Late Jurassic et al. 2013b; Benson et al. 2013; Benson and Bowdler 2014; (e.g., Bardet et al. 2014; Stubbs and Benton 2016; Foffa et Pérez-García 2015b, c; Püntener et al. 2015; Anquetin and al. 2018c; Zverkov et al. 2018; Fernández et al. 2019). Other Chapman 2016; Moon and Kirton 2018), the Reuchenette marine reptile clades, such as thalassochelydian turtles Formation (Kimmeridgian) of Switzerland (e.g., Rütimeyer and pleurosaurs (rhynchocephalians) were more restricted 1873; Bräm 1965; Meyer 1994; Comment et al. 2015; Pünte- in their geographic distribution and represented some of ner et al. 2015, 2017a, b; Sullivan and Joyce 2017; Raselli and the earliest-diverging, fully marine representatives within Anquetin 2019; Anquetin and Püntener 2020), the “Soln- Acta Palaeontol. Pol. 66 (1): 99–129, 2021 https://doi.org/10.4202/app.00795.2020 100 ACTA PALAEONTOLOGICA POLONICA 66 (1), 2021 hofen Limestone” (Tithonian) of Southern Ger many (e.g., ported from the Tithonian of Owadów-Brzezinki (Kin and Meyer 1839a; Parsons and Williams 1961; Gaffney 1975b; Błażejowski 2012; Kin et al. 2012, 2013). A partial skeleton Bardet and Fernández 2000; Dupret 2004; Young and de of an ichthyosaur was also reported, but not yet described, Andrade 2009; de Andrade et al. 2010; Young et al. 2012; from the Kimmeridgian of Szczerców (Grabowska 2013; Anquetin and Joyce 2014; Arratia et al. 2015; Anquetin et Maliszewski 2017; Krzeczyńska et al. 2020). Finally, several al. 2017; Bever and Norell 2017), several Kimmeridgian and teeth collected from Zalas (originaly reported by Molenda Tithonian localities of France (e.g., Thiollière 1850; Meyer 1997), Wapiennik, and Częstochowa (initially described by 1860; Lortet 1892; Rieppel 1980; Broin 1994; Lapparent de Maryańska 1972) were recently revised as belonging to tha- Broin et al. 1996; Bardet et al. 1997; Pérez-García 2015b) lassophonean pliosaurids (Lomax 2015; Tyborowski 2019). and Iberia (e.g., Lapparent de Broin and Murelaga 1999; A diverse assemblage of marine reptiles from the upper Pérez-García and Ortega 2011; Pérez-García 2015c), as well Kimmeridgian of Krzyżanowice was recently described by as several fossil-bearing horizons spanning the Volgian Tyborowski and Błażejowski (2019a, b) based on the revision (Tithonian–lowermost Berriasian) of European Russia (e.g., of historical specimens (Borsuk-Białynicka and Młynarski Zverkov et al. 2015a, b, 2018; Arkhangelsky et al. 2018; 1968) and the discovery of new fossil material. Although Zverkov and Efimov 2019; Zverkov and Prilepskaya 2019) both the historical and newly discovered specimens from and the Slottsmøya Member of the Aghardfjellet Formation Krzyżanowice are very fragmentary, Tyborowski and Bła- of Spitsbergen (e.g., Druckenmiller et al. 2012; Knutsen et żejowski (2019a, b) proposed that they represented a no- al. 2012a–c; Roberts et al. 2014, 2020; Delsett et al. 2019). table diversity of marine reptiles, including indeterminate Recent research has provided evidence for extensive “plesiochelyid” turtles, thalattosuchian crocodylomorphs faunal interchange between Late Jurassic seas located in (Machimosaurus sp.) and pliosaurid (Pliosaurus sp.) and Northern, Western and Eastern Europe (e.g., Arkhangelsky elasmosaurid (Elasmosauridae indet.) plesiosaurs. The tax- et al. 2018; Zverkov and Efimov 2019), with the seaway cov- onomic composition of this assemblage was suggested to ering Poland acting as an important dispersal route between be similar to that of the Kimmeridge Clay Formation of the them (Tyborowski 2016; Tyborowski and Błażejowski 2019a, United Kingdom and both assemblages were established as b). Therefore, marine reptile fossils from the Upper Jurassic part of the newly proposed “Matyja-Wierzbowski Line”—a of Poland have the potential for providing data important for transitional palaeobiogeographic region separating marine testing previously proposed hypotheses on the dispersal and faunas from Northern and Southern Europe (Tyborowski taxonomic diversification of European Late Jurassic ma- and Błażejowski 2019a, b). However, when reading the ac- rine reptiles. However, Late Jurassic marine reptile fossils counts of Tyborowski and Błażejowski (2019a, b) we identi- occur rarely in Poland and are often fragmentary or very fied the following problems regarding the taxonomic iden- incomplete (Fig. 1 and Table 1). This hinders their correct tification, provenance, and historical background provided taxonomic identification, crucial for the proper placement of for the fossils described therein: these fossils into biogeographic and evolutionary contexts. (i) The new material (MZ VIII Vr-71) referred to Plesio- Several localities yielding fragmentary fossils of Late chelyidae indet., which was described and figured by Tybo- Jurassic marine reptiles were reported from the territory rowski and Błażejowski (2019a, b) was determined to be of Poland during the 19th and 20th centuries (Fig. 1 and the same material previously described and figured by Table 1). However, the majority of these findings were only Młynarski and Borsuk-Białynicka (1968). The historical briefly described or merely mentioned in Polish or German nature of the material was not recognised by Tyborowski literature (e.g., Jentzsch 1884; Dames 1888; Gallinek 1895, and Błażejowski (2019a, b) and no attention was given to 1896; Deecke 1907; Hirszberg 1924; Molenda 1997; see the original, controversial designation of MZ VIII Vr-71 as Table 1) and have received little scientific attention until “Tretosternon aff. punctatum” Owen, 1842. very recently. Nevertheless, the last decade saw a marked (ii) The specimen
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